Cornisoane Pseudoantracnozice

There are few scientists of whom it can be said that their mistakes are more
interesting
than their colleagues' successes, but Albert Einstein was one. Few "blunders" have
had a
longer and more eventful life than the cosmological constant, sometimes described
as the
most famous fudge factor in the history of science, that Einstein added to his
theory of
general relativity in 1917. Its role was to provide a repulsive force in order to
keep the
universe from theoretically collapsing under its own weight. Einstein abandoned the
cosmological constant when the universe turned out to be expanding, but in
succeeding
years, the cosmological constant, like Rasputin, has stubbornly refused to die,
dragging
itself to the fore, whispering of deep enigmas and mysterious new forces in nature,
whenever cosmologists have run into trouble reconciling their observations of the
universe with their theories.
This year the cosmological constant has been propelled back into the news as an
explanation for the widely reported discovery, based on observations of distant
exploding
stars, that some kind of "funny energy" is apparently accelerating the expansion of
the
universe. "If the cosmological constant was good enough for Einstein," the
cosmologist
Michael Turner of the University of Chicago remarked at a meeting in April, "it
should
be good enough for us."
Einstein has been dead for 43 years. How did he and his 80-year-old fudge factor
come to
be at the center of a revolution in modern cosmology?
The story begins in Vienna with a mystical concept that Einstein called Mach's
principle.
Vienna was the intellectual redoubt of Ernst Mach (1838-1916), a physicist and
philosopher who bestrode European science like a Colossus. The scale by which
supersonic speeds are measured is named for him. His biggest legacy was
philosophical;
he maintained that all knowledge came from the senses, and campaigned relentlessly
against the introduction of what he considered metaphysical concepts in science,
atoms
for example.
1Another was the notion of absolute space, which formed the framework of Newton's
universe. Mach argued that we do not see "space," only the players in it. All our
knowledge of motion, he pointed out, was only relative to the "fixed stars." In his
books
and papers, he wondered if inertia, the tendency of an object to remain at rest or
in
motion until acted upon by an outside force, was similarly relative and derived
somehow
from an interaction with everything else in the universe.
"What would become of the law of inertia if the whole of the heavens began to move
and
stars swarmed in confusion?" he wrote in 1911. "Only in the case of a shattering of
the
universe do we learn that all bodies, each with its share, are of importance in the
law of
inertia."
Mach never ventured a guess as to how this mysterious interaction would work, but
Einstein, who admired Mach's incorrigible skepticism, was enamored of what he
sometimes called Mach's principle and sometimes called the relativity of inertia.
He
hoped to incorporate the concept in his new theory of general relativity, which he
completed in 1915. That theory describes how matter and energy distort or "curve"
the
geometry of space and time, producing the phenomenon called gravity.
In the language of general relativity, Mach's principle required that the space-
time
curvature should be determined solely by other matter or energy in the universe,
and not
any initial conditions or outside influences -- what physicists call boundary
conditions.
Among other things, Einstein took this to mean that it should be impossible to
solve his
equations for the case of a solitary object -- an atom or a star alone in the
universe --
since there would be nothing to compare it to or interact with.
So Einstein was surprised a few months after announcing his new theory, when Karl
Schwarzschild, a German astrophysicist serving at the front in World War I, sent
him just
such a solution, which described the gravitational field around a solitary star. "I
would
not have believed that the strict treatment of the point mass problem was so
simple,"
Einstein said.
Perhaps spurred in part by Schwarzschild's results, Einstein turned his energies in
the fall
of 1916 to inventing a universe with boundaries that would prevent a star from
escaping
its neighbors and drifting away into infinite un-Machian loneliness. He worked out
his
ideas in a correspondence with a Dutch astronomer, Willem de Sitter, which are to
be
published this summer by the Princeton University Press in Volume 8 of "The
Collected
Papers of Albert Einstein." Like most of his colleagues at the time, Einstein
considered
2the universe to consist of a cloud of stars, namely the Milky Way, surrounded by
vast
space. One of his ideas envisioned "distant masses" ringing the outskirts of the
Milky
Way like a fence. These masses would somehow curl up space and close it off.
His sparring partner de Sitter scoffed at that, arguing these "supernatural" masses
would
not be part of the visible universe. As such, they were no more palatable than
Newton's
old idea of absolute space, which was equally invisible and arbitrary.
In desperation and laid up with gall bladder trouble in February of 1917, Einstein
hit on
the idea of a universe without boundaries, in which space had been bent around to
meet
itself, like the surface of a sphere, by the matter within. "I have committed
another
suggestion with respect to gravitation which exposes me to the danger of being
confined
to the nut house," he confided to a friend.
This got rid of the need for boundaries -- the surface of a sphere has no boundary.
Such a
bubble universe would be defined solely by its matter and energy content, as
Machian
principles dictated. But there was a new problem; this universe was unstable, the
bubble
had to be either expanding or contracting. The Milky Way appeared to be neither
expanding nor contracting; its stars did not seem to be going anywhere in
particular.
Here was where the cosmological constant came in. Einstein made a little
mathematical
fix to his equations, adding "a cosmological term" that stabilized them and the
universe.
Physically, this new term, denoted by the Greek letter lambda, represented some
kind of
long range repulsive force, presumably that kept the cosmos from collapsing under
its
own weight.
Admittedly, Einstein acknowledged in his paper, the cosmological constant was "not
justified by our actual knowledge of gravitation," but it did not contradict
relativity,
either. The happy result was a static universe of the type nearly everybody
believed they
lived in and in which geometry was strictly determined by matter. "This is the core
of the
requirement of the relativity of inertia," Einstein explained to de Sitter. "To me,
as long
as this requirement had not been fulfilled, the goal of general relativity was not
yet
completely achieved. This only came about with the lambda term."
The joke, of course, is that Einstein did not need a static universe to have a
Machian one.
Michel Janssen, a Boston University physicist and Einstein scholar, pointed out,
"Einstein needed the constant not because of his philosophical predilections but
because
of his prejudice that the universe is static."
3Moreover, in seeking to save the universe for Mach, Einstein had destroyed Mach's
principle. "The cosmological term is radically anti-Machian, in the sense that it
ascribes
intrinsic properties (energy and pressure-density) to pure space, in the absence of
matter,"
said Frank Wilczek, a theorist at the Institute for Advanced Study in Princeton.
In any event, Einstein's new universe soon fell apart. In another 10 years the
astronomer
Edwin Hubble in California was showing that mysterious spiral nebulae were galaxies
far
far away and getting farther -- in short that the universe might be expanding.
De Sitter further confounded Einstein by coming up with his own solution to
Einstein's
equations that described a universe that had no matter in it at all.
"It would be unsatisfactory, in my opinion," Einstein grumbled, "if a world without
matter were possible."
De Sitter's empty universe was also supposed to be static, but that too proved to
be an
illusion. Calculations showed that when test particles were inserted into it, they
flew
away from each other. That was the last straw for Einstein. "If there is no quasi-
static
world," he said in 1922, "then away with the cosmological term."
In 1931, after a trip to the Mount Wilson observatory in Pasadena, Calif., to meet
Hubble,
Einstein turned his back on the cosmological constant for good, calling it
"theoretically
unsatisfactory anyway."
He never mentioned it again.
In the meantime, the equations for an expanding universe had been independently
discovered by Aleksandr Friedmann, a young Russian theorist, and by the Abbe
Georges
Lemaitre, a Belgian cleric and physicist. A year after his visit with Hubble,
Einstein
threw his weight, along with de Sitter, behind an expanding universe without a
cosmological constant.
But the cosmological constant lived on in the imagination of Lemaitre, who found
that by
judicious application of lambda he could construct universes that started out
expanding
slowly and then sped up, universes that started out fast and then slowed down, or
one that
even began expanding, paused, and then resumed again.
This last model beckoned briefly to some astronomers in the early 1950's, when
measurements of the cosmic expansion embarrassingly suggested that the universe was
4only two billion years old -- younger Earth. A group of astronomers visited
Einstein in
Princeton and suggested that resuscitating the cosmological constant could resolve
the
age discrepancy. Einstein turned them down, saying that the introduction of the
cosmological constant had been the biggest blunder of his life. George Gamow, one
of
the astronomers, reported the remark in his autobiography, "My World Line," and it
became part of the Einstein legend.
Einstein died three years later. In the years after his death, quantum mechanics,
the
strange set of rules that describe nature on the subatomic level (and Einstein's
bete noire)
transformed the cosmological constant and showed just how prescient Einstein had
been
in inventing it. The famous (and mystical in its own right) uncertainty principle
decreed
that there is no such thing as nothing, and even empty space can be thought of as
foaming
with energy.
The effects of this vacuum energy on atoms had been detected in the laboratory, as
early
as 1948, but no one thought to investigate its influence on the universe as a whole
until
1967, when a new crisis, an apparent proliferation of too-many quasars when the
universe
was about one-third its present size, led to renewed muttering about the
cosmological
constant. Jakob Zeldovich, a legendary Russian theorist who was a genius at
marrying
microphysics to the universe, realized that this quantum vacuum energy would enter
into
Einstein's equations exactly the same as the old cosmological constant.
The problem was that a naive straightforward calculation of these quantum
fluctuations
suggested that the vacuum energy in the universe should be about 118 orders of
magnitude (10 followed by 117 zeros) denser than the matter. In which case the
cosmological constant would either have crumpled the universe into a black hole in
the
first instant of its existence or immediately blown the cosmos so far apart that
not even
atoms would ever have formed. The fact that the universe had been sedately and
happily
expanding for 10 billion years or so, however, meant that any cosmological
constant, if it
existed at all, was modest.
Even making the most optimistic assumptions, Dr. Zeldovich still could not make the
predicted cosmological constant to come out to be less than a billion times the
observed
limit.
Ever since then, many particle theorists have simply assumed that for some as-yet-
unknown reason the cosmological constant is zero. In the era of superstrings and
ambitious theories of everything tracing history back to the first micro-micro
second of
unrecorded time, the cosmological constant has been a trapdoor in the basement of
5physics, suggesting that at some fundamental level something is being missed about
the
world. In an article in Reviews of Modern Physics in 1989, Steven Weinberg of the
University of Texas referred to the cosmological constant as "a veritable crisis,"
whose
solution would have a wide impact on physics and astronomy.
Things got even more interesting in the 1970's with the advent of the current crop
of
particle physics theories, which feature a shadowy entity known as the Higgs field,
which
permeates space and gives elementary particles their properties. Physicists presume
that
the energy density of the Higgs field today is zero, but in the past, when the
universe was
hotter, the Higgs energy could have been enormous and dominated the dynamics of the
universe. In fact, speculation that such an episode occurred a fraction of a second
after
the Big Bang, inflating the wrinkles out of the primeval chaos -- what Dr. Turner
calls
vacuum energy put to a good use -- has dominated cosmology in the last 15 years.
"We want to explain why the effective cosmological constant is small now, not why
it
was always small," Dr. Weinberg wrote in his review. In their efforts to provide an
explanation, theorists have been driven recently to talk about multiple universes
connected by space-time tunnels called wormholes, among other things.
The flavor of the crisis was best expressed, some years ago at an astrophysics
conference
by Dr. Wilczek. Summing up the discussions at the end of the meeting, he came at
last to
the cosmological constant. "Whereof one cannot speak, thereof one must be silent,"
he
said, quoting from Ludwig Wittgenstein's "Tractatus Logico-Philosophicus."
Now it seems that the astronomers have broken that silence.
Copyright 2002 The New York Times Company
6Mysteries of the Universe
Q U A N T U M P H Y S I C S
Quantum Theory Tugged, and All of Physics Unraveled
By DENNIS OVERBYE
They tried to talk Max Planck out of becoming a physicist, on the grounds that here
was
nothing left to discover. The young Planck didn't mind. A conservative youth from
the
south of Germany, a descendant of church rectors and professors, he was happy to
add to
the perfection of what was already known.
Instead, he destroyed it, by discovering what was in effect a loose thread that
when
tugged would eventually unravel the entire fabric of what had passed for reality.
As a new professor at the University of Berlin, Planck embarked in the fall of 1900
on a
mundane sounding calculation of the spectral characteristics of the glow from a
heated
object. Physicists had good reason to think the answer would elucidate the
relationship
between light and matter as well as give German industry a leg up in the electric
light
business. But the calculation had been plagued with difficulties.
Planck succeeded in finding the right formula, but at a cost, as he reported to the
German
Physical Society on Dec. 14. In what he called "an act of desperation," he had to
assume
that atoms could only emit energy in discrete amounts that he later called quanta
(from
the Latin quantus for "how much" ) rather than in the continuous waves prescribed
by
electromagnetic theory. Nature seemed to be acting like a fussy bank teller who
would
not make change, and would not accept it either.
That was the first shot in a revolution. Within a quarter of a century, the common
sense
laws of science had been overthrown. In their place was a bizarre set of rules
known as
quantum mechanics, in which causes were not guaranteed to be linked to effects; a
subatomic particle like an electron could be in two places at once, everywhere or
nowhere until someone measured it; and light could be a wave or a particle.
and the unknown, who as been accorded some of the ultimate accolades in pop culture
--
appearing as Einstein's poker buddy on "Star Trek: The Next Generation," and as a
guest
star on "The Simpsons."
While a graduate student, in 1963, he learned he had amyotrophic lateral sclerosis
and
was given a few years to live. He has moved about in a wheelchair for more than 25
years
and now speaks only through a voice synthesizer. Dr. Hawking, for whom the word
"puckish" seems to have been invented, has often said his disability is an
advantage
because it frees him to sit and think. Next month his colleagues will celebrate his
60th
birthday with a weeklong all-star symposium in Cambridge.
In the new book's introduction, Dr. Hawking admits that "A Brief History of Time"
was
"not easy going" and laments that some readers got stuck and did not finish it. He
has
tried, he says, to make this one easier. Slightly longer than the earlier book,
"Nutshell," at
216 pages, is embellished with colorful illustrations that give it a coffee-table-
book look.
So far the critics are in qualified agreement; one, Bryan Appleyard in the The New
he Bivalvia, the second largest class within the
Solnhofen Limestone of Eichsta tt, Germany, and was
described by Cosimo Collini (1727–1806) in 1784.
Collini concluded that it was a possible sea creature
of unknown affinity, although he did note bat-like
features. In 1801, the great French anatomist Georges
Cuvier (1769–1832) recognized that the creature
was a reptile and that its elongated digits must have
supported flight membranes. Cuvier was thus the first
to recognize pterosaurs as flying reptiles and, in 1809,
he coined the name ‘Ptero-Dactyle’. This later became
the generic name Pterodactylus (Figures 1 and 4).
In the decades that followed, a succession of further
pterosaurs from the Solnhofen Limestone was an-
nounced, many in a spectacular state of preservation
and some with their wing membranes intact. The
first recognized British pterosaur, a specimen of the
deep-skulled Dimorphodon, was discovered by Mary
Anning (1799–1847) in 1827 in Lower Jurassic rocks
of Lyme Regis, Dorset. We now know that Gideon
Mantell (1790–1852), best known for the discovery
of Iguanodon, found pterosaur remains before this
in the Early Cretaceous Wealden strata of Sussex,
but had thought that these were from birds. North
America yielded its first pterosaur to the prolific
palaeontologist O. C. Marsh (1831–1899) in 1871
and, by 1876, Marsh had recognized it as a new,
distinctive genus he named Pteranodon (meaning
‘winged and toothless’). With an estimated wingspan
of 6 m, Pteranodon was huge compared to most
earlier discoveries.
While these discoveries and others were being
made, varied opinions on the nature and life style of
pterosaurs were appearing, and they were variously
depicted as swimming creatures, as bats, marsupials,
or as kin of birds. By the early 1900s, it was generally
agreed that pterosaurs were bat-like flying reptiles and,
in 1901, Harry Seeley (1839–1909) published Dragons
of the Air, the first book devoted to pterosaurs.
South American Cretaceous pterosaurs have proved
to be among the most important in the world, but not
until 1971 was the first pterosaur from the now famous
Santana Formation of Brazil discovered. Since then a
significant number of new kinds from around the
world (around 70 genera are presently recognized)
have revealed previously unimagined morphologies
and maximum sizes. Until 1971, Pteranodon sternbergi
Figure 1 Life restoration of the Late Jurassic pterodactyloid, Pterodactylus , from
the German Solnhofen Limestone in a quadrupedal
stance. Note the presence of body hair and the soft tissue head crest. Reproduced
with permission from Dino Frey. Buffetaut E
and Mazin J M (2003) Evolution and Palaeobiology of Pterosaurs. Geological Society
Special Publication 217 . London: The Geological Society
of London.510 FOSSIL VERTEBRATES/Flying Reptiles
(wingspan, 9 m) was the largest known flying animal,
but the discovery in Texas of Quetzalcoatlus revealed
that the biggest pterosaurs achieved wingspans of
11 m. Related pterosaurs of similar or larger size were
discovered in the 1990s in Spain and eastern Europe.
The Pterosaur Skeleton
The pterosaur skeleton was highly modified for flight,
and the most obvious features are the huge size of the
skull compared with the body and the extreme
elongation of one of the fingers. Like birds, most
pterosaurs had hollow bones with foramina (small
openings), indicating that they contained air sacs
connected to the lungs. Pterosaur bones were sup-
ported internally by struts, and the bone walls them-
selves, usually no thicker than 2 mm, are composed
of multiple overlapping layers and thus combine
lightness with strength.
Pterosaur skull morphology is varied, although the
majority had long, slim, shallow jaws and all had
large orbits (eye-sockets). In basal pterosaurs, the
external nostril was separate from an opening in
front of the orbit called the antorbital fenestra. In
pterodactyloids, these two openings merged into a
single one called the nasoantorbital fenestra. Ptero-
saur teeth were extremely variable. Widely spaced
pointed teeth, were widespread and from ancestors
with teeth like these evolved species with fang-like
teeth at the jaw tips and the unique Istiodactylus
with its short petal-shaped teeth. The Late Triassic
Eudimorphodon and Austriadactylus possessed
multicusped teeth while elongate, slender teeth
numbering in the hundreds evolved in the ctenochas-
matoids. Toothlessness evolved several times. Some
pterosaurs skulls sport bony crests at the jaw tips,
along the midline or at the back of the skull.
Figure 2 Wing skeleton of an ornithocheiroid pterodactyloid.
Unlike birds and bats, the main wing spar in ptero-
saurs was formed by a hypertrophied digit (Figure 2).
This ‘wing finger’ is generally considered to be the
fourth because the digital formulae of the pterosaur
hand best matches that of digits one to four in the
hands of other reptiles. However, a rod-shaped bone
projecting from the pterosaur wrist, called the pteroid
bone, has at times been argued to represent the first
hand digit. This is a minority view today but, if it is
correct, then pterosaurs have five hand digits and the
wing finger is the fifth. Although most pterosaur
fossils show the pteroid pointing towards the shoul-
der, some workers suggest that it pointed forwards
parallel to the neck. Regardless, the pteroid was
probably mobile and used to control the attitude of
the propatagium (Figure 3).
The pterosaur pectoral girdle includes a (normally
fused) scapula and coracoid that meet at an acute
angle and, as expected for flying animals, the socket
for the humerus faces sideways and slightly upwards.
The coracoids attach to an enlarged keeled sternum
that anchored most of the major flight muscles. The
bones of the pterosaur pelvis were short, usually fused
together, and with a closed hip socket. Pterosaurs
have a pair of unique rod- or plate-like bones called
the prepubes projecting forwards from the bottom
of the pelvis. Like the gastralia (‘belly ribs’) that all
pterosaurs possessed, they may have helped support
the gut or keep the abdomen rigid.
The vertebral column in pterosaurs can clearly be
differentiated into cervical, dorsal, sacral, and caudal
portions. The number of vertebrae is variable and
pterosaurs have 7–9 cervical, 11–16 dorsal, 3–10
sacral, and 11–40 caudal vertebrae. The many caudal
vertebrae of basal pterosaurs are encased in long bony
processes that make the tail stiff and rod-like. In
derived Cretaceous pterosaurs, most of the dorsalFOSSIL VERTEBRATES/Flying Reptiles
511
vertebrae are fused together forming a structure
called the notarium.
Compared with the other wing segments, the ptero-
saur humerus is short, although generally with mas-
sive crests for muscle attachment. The ulna is always
larger than the radius and both are attached distally
to block-shaped carpal bones. Projecting from one
of these is the unique pteroid. Pterosaurs had four
metacarpals, the first three of which were slim and,
with the exception of Nyctosaurus from Late Cret-
aceous North America, attached to short, clawed
fingers. Why Nyctosaurus lacked clawed fingers is
unknown, but in all other pterosaurs these digits
may have served important functions. Trackways
show that they were used in walking, and it is also
possible that they were employed in grooming or
climbing. The fourth metacarpal was robust and
tipped with a twisted, roller-like distal end to which
was attached the massive wing finger. This consists of
four long straight bones, excepting a few genera
where there were only three. Because of the twisted
end of the fourth metacarpal, the wing finger would
have lain parallel to the body’s long axis when the
wing was folded up.
The pterosaur hind limb is lightly built and the
head of the femur is only slightly offset from the
long axis of the shaft. Pterosaur hind limbs seem to
have been quite flexible, but mostly sprawled to the
sides. During flight, the hind limb was probably held
in a bat-like orientation and could have been used to
control the shape of the wing membranes. Basal
pterosaurs are five-toed, with a prominent curving
Figure 3 Schematic representation of the flight membranes in
a generalized pterodactyloid pterosaur.
fifth digit that is hooked towards the tail. In ptero-
dactyloids, the fifth digit is either absent or present as
a tiny stub.
Because some articulated fossils indicate that the
foot could assume a 90 # angle relative to the tibia
(and there is little evidence for much motion at the
metatarsophalangeal joints), pterosaurs have gener-
ally been regarded as plantigrade (placing the whole
length of the foot on the ground when walking). In
1983, Kevin Padian argued that this was not the case
for Dimorphodon and that it may instead have been
digitigrade (walking only on the toes). This was later
inferred for all pterosaurs. An articulated Dimorpho-
don foot shows, however, that only limited motion
was possible at the metatarsophalangeal joint, thus
supporting a plantigrade posture. This is in agreement
with probable pterosaur tracks preserved as trace
fossils.
Soft Tissue, Integument, and
Pterosaur Life Appearance
Many aspects of pterosaur life appearance remain
unknown or controversial, although a number of ex-
ceptional fossils have provided some surprising
details. Pterosaur body hair was reported as early as
1831 and described for various Jurassic pterosaurs
between the 1920s and 1970s and today it is clear
that pterosaurs had bristle-like hairs covering their
necks and bodies (Figure 4). The active flapping flight
and body hair of pterosaurs suggest that they had an
elevated metabolism.
Other exceptional fossils show that some ptero-
saurs possessed a throat pouch, webbing between
the toes, and scales on the soles of the feet. Soft
Figure 4 An exceptionally well preserved skeleton of the Late
Jurassic pterodactyloid Pterodactylus from the German Solnhofen
Limestone. This specimen preserves parts of the flight mem
branes, a throat pouch, and hairs on the neck and back.512 FOSSIL
VERTEBRATES/Flying Reptiles
Figure 5 Variation in skull crest morphology in pterodactyloids. Soft tissue crests
are now known for a wide diversity of
pterodactyloids. Reproduced with permission from Dino Frey and Marie Celine Buchy.
Buffetaut E and Mazin J M (2003) Evolution
and Palaeobiology of Pterosaurs. Geological Society Special Publication 217 .
London: The Geological Society of London.
tissue skull crests connected to the underlying bony
crests have proved to be widespread and appear
to have doubled the size of the bony crests (Figures 5
and 6). An unexpected discovery is a soft tissue crest
in Pterodactylus, a genus that lacks a bony crest
(Figure 1). The presence of a distinctive bone texture
on the pterosaur snout, jaw, and palate indicates that
pterosaurs were beaked.
Pterosaur wing membranes are known from well-
preserved specimens from the Solnhofen Limestone
and the Early Cretaceous Brazilian Crato and San-
tana formations. A membrane called the propatagium
extended from the shoulder to the pteroid and per-
haps distally to encompass the first three fingers. The
main flight membrane, the brachiopatagium (also
called the cheiropatagium), extended from the tip of
the wing finger to the hind limb, extending
as far distally as the knee, shin, or ankle. Another
membrane, the uropatagium, was present between
the hind limbs (Figure 3). The wing membrane
appears to have been complex, with a thin epidermis,
a layer of vascular tissues, a layer of stiffening fibres
called aktinofibrils, a thin sheet of muscle, and a
Figure 6 Skull of the tapejarid pterosaur Topejara navigaus from
the Early Cretaceous Crato Formation of Brazil with bony and soft
tissue skull crest. Buffetaut E and Mazin J M (2003) Evolution and
Palaeobiology of Pterosaurs. Geological Society Special Publication 217 .
London: The Geological Society of London.FOSSIL VERTEBRATES/Flying Reptiles 513
blood capillary network. Rhamphorhynchus and
probably other long-tailed pterosaurs possessed a
vertical diamond-shaped membrane at the tail tip.
With skin membranes connecting the wings, body,
and legs, pterosaurs may have been superficially bat-
like but, because bats are mostly dark-coloured noctur-
nal animals, it is doubtful that the similarities were
strong. Pterosaurs mostly seem to have been ecological
analogues of sea- and water-birds, and it might be
that they were patterned in whites, blacks, and greys,
although bright colours presumably decorated their
crests.
pterosaur hind limbs are only superficially similar to
those of dinosaurs, and that re-analysis favoured a
position for pterosaurs outside of the crocodilian–
dinosaur group. Rather more heterodox recent ideas
include the suggestion that pterosaurs are the closest
relatives of birds and that pterosaurs are part of the
Dinosauria.
Several different models have been proposed for
the origin of pterosaurs, but the presence in basal ptero-
saurs of climbing features and of various details in the
hind limb and pelvis indicative of a leaping ability sug-
gest that pterosaurs first evolved as tree-climbing
leapers.
The Affinities and Origin of Pterosaurs
Historically, pterosaurs have been allied with Meso-
zoic marine reptiles, bats, marsupials, and birds
(see Fossil Vertebrates: Dinosaurs; Birds). However,
major improvements in the understanding of verte-
brate evolution allowed the palaeontologists of the
nineteenth and twentieth centuries to realize that
pterosaurs were related at least vaguely to dinosaurs
(see ) and their allies.
Although it is clear that pterosaurs are part of the
major reptile assemblage known as the Diapsida,
their affinities within this group are controversial.
The presence of an antorbital fenestra has conven-
tionally meant that pterosaurs have been regarded
as archosaurs, the so-called ruling reptile group
that incorporates crocodilians, dinosaurs, and kin.
Among archosaurs, pterosaurs share a simple hinge-
like ankle joint with dinosaurs and consequently
have been regarded as close relatives of dinosaurs in
most studies. This view was developed at a time when
some workers thought that pterosaurs originated
from terrestrial bipedal ancestors and that pterosaurs
themselves were bipedal and digitigrade. A small bi-
pedal, long-legged archosaur from Late Triassic
Scotland, Scleromochlus, was argued to be a ptero-
saur ancestor, but recent studies refute this idea. The
idea that pterosaurs might be close relatives of dino-
saurs can certainly be regarded as the ‘mainstream’
view in vertebrate palaeontology today. However,
several recent studies have questioned the evidence
for this proposed affinity.
An alternative hypothesis argues that pterosaurs
belong instead to a group of archosaur-like diapsids,
the Prolacertiformes. Most prolacertiforms were
superficially lizard-like, but Sharovipteryx from Late
Triassic Kyrgyzstan appears to be intermediate be-
tween conventional prolacertiforms and pterosaurs.
It has pterosaur-like hind limbs and vertebrae and
membranes between its hind limbs and tail.
Some other models for pterosaur ancestry have been
proposed. In 1996, S. Christopher Bennett argued that
Pterosaur Diversity and Phylogeny
It was recognized in 1901 that pterosaurs could be
divided into two groups: the toothed, mostly long-
tailed Rhamphorhynchoidea, and the short-tailed
Pterodactyloidea (including both toothed and tooth-
less kinds). Today, it is clear that rhamphorhynchoids
include the ancestors of pterodactyloids and, conse-
quently, Rhamphorhynchoidea is a grade and not a
clade. Pterosaurs previously referred to as rhamphor-
hynchoids are nowadays termed basal pterosaurs or
non-pterodactyloids. Although basal pterosaurs were
diverse, it is notable that they were small compared
with the majority of Cretaceous pterodactyloids.
The evolutionary relationships of pterosaurs are
relatively understudied and only recently has pterosaur
phylogeny been analysed. Although some areas of con-
sensus have emerged, authors disagree on the details.
We follow the phylogeny proposed by David Unwin
of the Museum fu r Naturkunde in Berlin (Figure 7).
Perhaps the most basal pterosaur is Preondactylus
from the Late Triassic of Italy. This form has a shorter
coracoid and humerus and longer legs than other
pterosaurs. Dimorphodontids, which include Dimor-
phodon from Early (and perhaps Middle) Jurassic
Figure 7 Cladogram depicting the relationships of all the major
pterosaur groups. Reproduced from David Unwin.514 FOSSIL VERTEBRATES/Flying
Reptiles
England and Mexico, are basal pterosaurs with deep
skulls superficially like those of puffins, whilst anur-
ognathids were unusual in having short, broad snouts
and abbreviated tails. A surprising recent discovery
is the persistence of anurognathids into the Early
Cretaceous. Two basal pterosaurs, Eudimorphodon
from Late Triassic Italy and Greenland and Campy-
lognathoides from Early Jurassic Germany and India,
are united in the Campylognathoididae based on a
distinctive lower jaw in which two pairs of large
conical teeth are followed by multiple smaller ones.
Rhamphorhynchids were successful Jurassic ptero-
saurs known from Eurasia, North America, and
Africa. Rhamphorhynchus from Late Jurassic Europe
exhibits a laterally compressed, ventrally directed
lower jaw tip and an array of forward-pointing
teeth. It probably used these to grab fish and other
small prey from the water. Another rhamphorhynchid
lineage, the scaphognathines, had deeper skulls with
teeth perpendicular to the jaw margins.
The Pterodactyloids
Pterodactyloids, the advanced short-tailed pterosaurs,
originated from a rhamphorhynchid-like ancestor
during the Middle Jurassic. The pterodactyloid radi-
ation consisted of four major groups: the robust-jawed
ornithocheiroids, the slim-jawed ctenochasmatoids,
the low-crested dsungaripteroids, and the long-necked,
crested azhdarchoids. A fifth group, the lonchodectids
from Early Cretaceous England, are of uncertain affin-
ity. Lonchodectids were small (wingspan, 1–2 m) with
long, dorsoventrally flattened jaws with small teeth,
each of which was supported by a low bony collar at
its base.
Ornithocheiroids were large predatory pterosaurs
(wingspan, 2–9 m) with robust beaks, often housing
recurved, fang-like teeth at their tips (Figure 8). Their
jaws frequently possessed keel-like dorsal and ventral
crests, and some forms also possessed crests on the
back of the skull. The toothless pteranodontids and
nyctosaurids appear to be members of this group. The
earliest known ornithocheiroids appear at the start of
the Cretaceous, while nyctosaurids survived to the
very end of the Cretaceous.
Ctenochasmatoids had needle-like meshes of teeth
set in long, thin jaws. In Pterodaustro from late
Early Cretaceous Argentina, the upturned lower jaw
contains approximately 1000 bristle-like teeth. These
were surely used for filtering small organisms from
the water. Unlike ornithocheiroids, ctenochasmatoids
had elongate cervical vertebrae and were generally
small (wingspan, 50 cm to 2 m), although Cearadac-
tylus from Early Cretaceous Brazil was a giant with
a wingspan of 5.5 m.
Figure 8 Jaw tips of the Cretaceous ornithocheiroid pterosaur
Coloborhynchus . Note the massive fang like anterior teeth and the
low keel like crests on both jaws. This specimen is from the
Santana Formation of Brazil and would have belonged to an
animal with a skull of approximately 1 m in length.
Members of the Dsungaripteroidea are known from
the Late Jurassic and Early Cretaceous of Eurasia,
Africa, and South America. Like some ctenochasma-
toids, dsungaripteroids had a midline crest on the top of
the skull. Their beaks often had toothless tips and they
may have been predators of molluscs and crustaceans.
Finally, the strangest pterodactyloids must be the
azhdarchoids. These include the long-necked azh-
darchids and the crested tapejarids. Azhdarchids
may have exceeded wingspans of 11 m and were
widely distributed in the Late Cretaceous. They may
have been ecological generalists akin to storks, and
were probably not specialist carrion feeders or mud
probers as has been proposed. Determining the life
style of the tapejarids is more difficult. The vaguely
parrot-like skull of Tapejara (Figure 6) from Lower
Cretaceous Brazil led some workers to propose that
it was a fruit eater, but it might better be imagined
as an auk analogue. Recently, it has been suggested
that Thalassodromeus, also from Lower Cretaceous
Brazil, was a fish eater that trawled its blade-like
lower jaw through the water.
Pterosaur Palaeobiology
Because pterosaurs are unique and extinct, recon-
structing their palaeobiology is difficult and nothing
is known about several aspects of their lives. Limited
evidence does allow us, however, to reconstruct their
sensory abilities, feeding behaviours, and styles of
locomotion.
The large orbits of pterosaurs show that they had
large eyes, and the abundance of visual display
phylum Mollusca, is one of the most familiar of all
invertebrate taxa. Modern representatives, such as
mussels, cockles, oysters, and scallops, are well
known from excursions to the coast, and in many
parts of the world they are important commercial
species. Their generally excellent fossil record has
allowed their evolutionary history to be traced back
to the Early Palaeozoic and, for much of this time,
they have been important components of many
faunas. From rather modest beginnings, they have
conquered a range of habitats from the deep sea to
freshwater, exploited a wide range of life habits
(from deep burrowing to swimming), and undergone
a near-exponential taxonomic proliferation, a spec-
tacular example of an adaptive radiation.
Bivalves come in all manner of shapes and sizes,
from tiny, thin-shelled commensals that live in as-
sociation with sea anemones, to giant clams and the
extinct rudists and inoceramids which reach(ed)
sizes well over 1 m. Shell morphology is extremely
plastic, but all are modifications of the same basic
theme. The intimacy of the shell morphology to life
habit has been a great benefit in reconstructing the life
habits of extinct bivalves, but has also frustrated
many attempts to establish the relationships between
different groups within the class. Bivalves have been
proven to be good palaeoenvironmental indicators,
but they have only limited use in biostratigraphy.
Freshwater ‘mussels’ have been used to date fluvial
deposits in the Carboniferous Coal Measures of
Western Europe, and inoceramids have been used
for Late Cretaceous deep marine settings (e.g., in
New Zealand). In general, however, species of the370 FOSSIL INVERTEBRATES/Bivalves
class are too long lived and too facies specific to be of
any great value.
General Morphology
As the name implies, bivalves comprise two calcar-
eous valves. These are arranged laterally (left and
right), are joined dorsally by a partially calcified elas-
tic ligament, and enclose the soft tissue. Each valve
has clearly differentiated posterior and anterior fea-
tures, i.e., inequilateral. The primitive arrangement,
retained by most bivalves, was to have a plane of
symmetry parallel to the commissure (the join be-
tween the two valves), resulting in valves which are
mirror images of one another (i.e., equivalve). Al-
though this symmetry is found in virtually all bivalves
which live with the commissural valve perpendicular
to the substrate surface (orthothetic), it has been lost
in those which have adopted a pleurothetic habit
where they lie on one valve (e.g., oysters, scallops).
In these cases, there is a tendency for the two valves to
become dissimilar (i.e., inequivalve), typically with
the underlying valve becoming more bowl-like and
the ‘upper’ one more reduced like a lid.
Shell Morphology
All bivalves possess a pair of shells which may be
shut to provide protection from both environmental
stresses (e.g., desiccation in the intertidal habitat) and
the threat of predation. Most shells are reasonably
robust, which has provided the class with a generally
excellent fossil record. Although shell morphology in
bivalves is very variable and intimately linked to their
life habits (see below), all shells are simple modifica-
tions of the basic shell secretion model used by all
shelled molluscs. The shell is secreted by the mantle
lobes and grows by marginal accretion, as evidenced
by the growth lines on the surface of the valve
(Figures 1A and 2A). These growth lines are particu-
larly marked in bivalves from intertidal and shallow
temperate habitats, where the animals experience
pronounced seasonality and largely stop growing
during the winter months. Bivalves which experience
more equable conditions do not show such obvious or
regular patterns. Inspection of the growth lines in
sectioned valves shows that, although most shell ma-
terial is added ventrally, the shell is also thickened
during growth (Figure 2B), demonstrating that the
entire mantle surface is responsible for adding mater-
ial. The outermost part of the shell is an organic layer
called the periostracum secreted at the mantle edge
(Figure 2C). The thickness of the periostracum varies
between taxa, from less than 1 mm in oysters and
Figure 1 (A) Mercenaria mercenaria , a shallow burrowing bi
valve from the Pliocene of Florida. Note the prominent annual
growth bands. (B) Pecten maxima , a free living epifaunal scallop
from the Holocene Atlantic.
scallops to several hundred micrometres in some
mussels. In many cases, the periostracum is lost by
abrasion and decay during the life of the animal,
particularly on the older parts of the shell, and there
is no real prospect of it being preserved in any but the
most exceptional circumstances. The primary func-
tion of the periostracum is to act as the template on
which the calcareous part of the shell is deposited, but
it may also provide protection from both corrosive
waters and predators that dissolve the shell. It is
particularly noticeable that freshwater bivalves have
very thick periostraca.
The main part of the shell, however, is calcareous.
It is in effect a ceramic made up of calcium carbonate
crystals in an organic matrix (the latter accounting for
<5% of the dry weight of the shell). The protein-
aceous matrix controls both the polymorph of cal-
cium carbonate used and the arrangement of the
crystals. All bivalves contain aragonite in their shells
and the vast majority are wholly so. Some taxa, how-
ever, chiefly those exploiting epifaunal life habits, also
secrete calcite in their outer layers. The oysters have
taken this to its extreme and the bulk of the shell isFOSSIL INVERTEBRATES/Bivalves
371
Figure 2 Details of shell formation in a generalized bivalve.
(A) Marked comarginal growth lines on the shell surface.
(B) Section through the shell along the line indicated in (A) show
ing the arrangement of growth lines within the shell. (C) The
relationship between the shell and the underlying mantle edge
(close up details of the circled area in (B)). i, inner mantle fold; m,
middle mantle fold; o, outer mantle fold.
calcitic, with aragonite being confined to the sites of
muscle attachment and the ligament.
Molluscan shell is immensely strong, in fact often
much stronger than vertebrate bone. There are a
number of different microstructures (Figure 3), each
with different mechanical properties, and most shells
are made up of two or three arranged in different
layers. Different taxa show different arrangements
and these are considered to be of phylogenetic signifi-
cance. It is apparent that the earliest bivalves were
wholly aragonitic and chiefly composed of nacre
(Figure 3A), and that subsequent evolution has pro-
duced the wide array of microstructural arrangements
seen today. The effect of differing crystal sizes, amount
of organic material, and polymorph used has affected
the preservation potential of different taxa; many of
the Palaeozoic and Mesozoic taxa that were originally
aragonitic are either preserved as internal moulds or
are replaced by calcite.
Details of the internal features of the shell are
shown in Figure 4. The hinge plate is situated dorsally
and houses the ligament and teeth. The ligament is an
elastic, partially calcified layer that provides a very
energy-efficient opening mechanism. During valve
closure, energy is stored in the ligament as it is flexed
by the contracted adductor muscle(s) (Figure 4C).
When the muscle is relaxed, the ligament springs the
valves apart causing them to gape. This passive valve
opening mechanism is the reason why many fossil
bivalves are found in a disarticulated state. Although
the ligament itself is seldom preserved, its position
may be inferred from the presence of the ligament
pits in which it is anchored (Figures 4 and 5). Most
bivalves have teeth on the hinge plate which fit into
corresponding sockets on the opposite valve and func-
tion to keep the valves in perfect alignment. Both
ligamenture and dentition vary markedly amongst
higher taxa of bivalves, and both are often used as
informative characters in establishing phylogenies.
Some of the range of hinge plate architectures is
shown in Figure 5.
A number of attachment scars mark the locations
where muscles are anchored to the shell. The most
significant of these are the adductor scars (Figures 4A
and 4B). If the adductor scars are paired (i.e., dimyar-
ian), they occur posteriorly and anteriorly. If an
animal is monomyarian, the single muscle (the pos-
terior) occupies a more central position. In many
taxa, there is a thin pallial line running around the
shell a small distance from the ventral edges that
marks the attachment of the mantle to the shell. In
infaunal taxa, where the posterior mantle has been
fused and elongated to form siphons, the pallial line is
inflected forming the pallial sinus. The sinus repre-
sents the space into which the siphons are withdrawn
when the valves are shut. Other muscle attachment
scars may be more or less apparent, including the
insertions of the pedal musculature (particularly in
burrowers and byssate taxa).
Soft Part Anatomy
Bivalves are laterally compressed and, unlike most
molluscs, there is no head or radula. The internal
organs are enclosed by the two mantle lobes that are
joined dorsally (Figure 4C). The chief function of the
mantle is to secrete the shells, but the ventral edges of
each mantle lobe are differentiated into three folds
(Figure 2C), only the outermost of which is directly
concerned with shell manufacture. The innermost
fold controls water flow into and out of the mantle
cavity, whilst the middle fold has sensory capability.
In several bivalve groups (such as scallops), the
middle fold is well developed with tentacles and
eyes. In some taxa, the mantle is extended posteriorly
and fused to form a pair of siphons through which
water is directed into (inhalant) and out of (exhalant)
the mantle cavity.372 FOSSIL INVERTEBRATES/Bivalves
Figure 3 Scanning electron micrographs of four of the most common bivalve shell
microstructures. (A) Aragonitic nacre (from the
inner shell layer of Pinna nobilis ). (B) Aragonitic crossed lamellar structure
(from the outer layers of Corbula gibba ). (C) Foliated calcite
(from Ostrea edulis ). (D) Calcitic prisms (from the outer shell layer of Pinna
nobilis ). Scale bars for (A) (C) represent 10 mm and for
(D) represents 100 mm.
The mantle cavity is spanned by one or two adductor
muscles that attach to the shell and act antagonistically
with the ligament to close the valves on contraction
(Figure 4C). A significant part of the mantle cavity is
occupied by a pair of gills (the ctenidia) lying on either
side of the rest of the viscera. In most bivalves, the gills
are involved with both respiration and ciliary suspen-
sion feeding (filtering small particles out of the water
which are then transferred to the mouth by a pair of
labial palps). Recent bivalves show a number of differ-
ent gill morphologies depending largely on the feed-
ing process employed. Deposit feeders, e.g., Nucula,
have less well-developed (protobranch) gills, whilst
members of the Lucinidae augment their filter feeding
by energy gained from the activities of sulphide-oxidiz-
ing chemosymbiotic bacteria living within the modified
gills. The carnivorous septibranch bivalves (e.g.,
Cuspidaria, Poromya) suck in small prey (such as
amphipods) using their modified siphons. These ex-
traordinary bivalves have ‘lost’ their gills and respire
over the inner surface of the mantle. Other significant
organs within the mantle cavity include the gut, heart,
circulatory system, and the foot. The gut runs between
the anteriorly positioned mouth and the posterior anus,
and includes a complex stomach which, again, has a
number of configurations depending on the feeding
biology of the animal. Blood is circulated throughout
the animal by a three-chambered heart. A muscular
foot is present in all juvenile and most adult bivalves
and occupies the centre of the mantle cavity.
Naturally, the soft part anatomy of bivalves is very
seldom preserved, although preservation of gill and
muscle material has been reported in exceptional cir-
cumstances. Various details, however, can be inferred
from the study of the internal surface of the shells.
Apart from adductor muscle scars, the practised eye
may pick out the attachment points of more minor
muscles and impressions of radial muscles and blood
vessels within the mantle.
Ecology
Modern bivalves exploit a wide range of life habits.
Many burrow to varying depths within soft sediments,FOSSIL INVERTEBRATES/Bivalves
373
Figure 4 Various aspects of the internal morphology of bivalve
shells. (A) The left valve of a generalized burrowing dimyarian.
(B) The right valve of a generalized byssate scallop. (C) The rela
tionship between the two valves and their associated mantle
lobes, adductor musculature and ligament.
but others attach to or bore into hard substrates by a
variety of means; others have become free living, some
with the ability to swim. Most bivalves are marine,
exploiting niches from the intertidal zone down into
the abyssal depths, but successful groups (including
the oysters) have invaded more brackish conditions
and even freshwater, where modern unionid mussels
cause enormous damage as biofoulers. It is clear that
the most primitive bivalves were marine shallow
burrowers and that other life styles evolved later. It is
also apparent that many of the more specialized life
habits have evolved separately in a number of differ-
ent lineages (i.e., polyphyletically). Seminal work by
S. M. Stanley firmly established how different aspects
of the morphology of living bivalves could be related
to their life habits, such that it is possible to use these
characteristics of extinct taxa to reconstruct the life
habits of fossil groups.
Burrowing
A large proportion of all bivalves (around 50% of all
modern families) burrow into soft sediments using the
foot. Most are equivalve and are isomyarian (i.e., the
posterior and anterior adductor muscles are of equal
size). The depth to which they burrow varies between
taxa, from those which lie just under the surface with
the edge of the shell virtually level with the sediment–
water interface (e.g., Cerastoderma; Figure 6C), to
depths of several centimetres (e.g., Mya; Figure 6B),
with Panopea reaching spectacular depths of up to
1 m. The key to successful burrowing is maintaining
contact with the seawater in order to continue
both feeding and respiration. This is achieved by
the siphons, snorkel-like extensions of the posterior
mantle. The length of the siphons, and therefore the
depth of burrowing, can be inferred from the shells by
the size of the pallial sinus; deeper burrowers have
more indented pallial sinuses, whereas very shallow
burrowers have no sinus at all (see Figure 6). Very
deep burrowers, such as Mya, have siphons so long
that they are unable to withdraw them fully into the
shell when it shuts, and have a permanent posterior
siphonal gape through which they protrude. Shallow
burrowers generally have strong, robust shells, often
with a pronounced radial or concentric ornament that
may assist the burrowing process or help the animal
remain ‘locked’ into the sediment. Deeper burrowers
tend to have thinner shells and are often smooth
shelled. Although the foot is never preserved, its pres-
ence may be inferred from the pedal musculature on
the inside of the valves and, in cases where the animal
is a rapid and deep burrower (such as the razor shell
Ensis), the foot may be so well developed as to require
an anterior pedal gape.
It is clear from studies of the siphons of living
bivalves that they are constructed in a number of dif-
ferent ways, suggesting that the deep burrowing habit
has evolved independently in several clades.
Attachment
Almost all larval bivalves attach to the substrate, if
only briefly, with tanned protein threads (the byssus)
secreted by a gland at the base of the foot. In a large
number of taxa, this habit has been neotenously
retained into adulthood, and again it is clear that
this has happened repeatedly in different groups.
Byssate bivalves fall into two categories: those like
Pinna (Figure 6A) and Modiolus that are orthothetic
and live attached to clasts within the sediment in
which they are partially buried (endobyssate), and
those that are attached to the surface of hard
substrates (epibyssate), either in an orthothetic (e.g.,
Mytilus; Figure 7D) or a pleurothetic (e.g., Isogno-
mon ephippium; Figure 7C) orientation. Orthothetic
byssate bivalves tend to be equivalve and have much
reduced anteriors. This anterior reduction is reflected374 FOSSIL
INVERTEBRATES/Bivalves
Figure 5 Selected hinge plates showing some of the variety of ligament insertion
and arrangement of teeth. (A) Cerastoderma edule :
heterodont dentition with two centrally placed cardinal teeth and two lateral
teeth. (B) Venus casina : heterodont (similar to Cerastoderma
but with no lateral teeth). (C) Arca tetragona : taxodont dentition with numerous
teeth arranged in a row; the ligament forms a chevron
pattern on the broad triangular area below the umbones. (D) Chlamys varia : two
simple teeth with the internal ligament occupying a
triangular pit below the umbones.
Figure 6 Morphology and mode of life of bivalves living in or partially within soft
substrates. (A) Pinna nobilis . Not to scale. (B) Mya
truncata . (C) Cerastoderma edula .
in the adductor musculature, which (although still
dimyarian) is heteromyarian, with the anterior ad-
ductor much smaller than the posterior (Figure 7D).
Pleurothetic byssate bivalves are often markedly
inequivalve, with the ‘lower’ valve (which in the ma-
jority of cases is the right) often larger than the other.
Although they are dimyarian early in ontogeny, the
‘adults’ are monomyarian, having lost the anterior
muscle during ontogeny; the remaining posterior
muscle is often large and centrally placed (Figure 7C).
The presence of a byssus may be inferred from either a
slight gape between the valves through which it passes
(the byssal gape), or more obviously the byssal notch in
scallops (Figure 4B).FOSSIL INVERTEBRATES/Bivalves 375
Figure 7 Morphology and mode of life of bivalves living on or boring into
substrates. (A) Spengleria rostrata . (B) Spondylus americanus .
(C) Isognomon ephippium . (D) Mytilus edulis . (E) Placuna placenta . (F) Gryphaea
arcuata . Not to scale.
Whereas byssate attachment is flexible and also
renewable, some bivalves permanently attach to
hard substrata by a calcareous cement. The cemented
habit always succeeds a byssate phase and it is clear
that it has evolved independently in a number of dif-
ferent clades (e.g., oysters, rudists, and a number of
pectinoids including Spondylus). Cemented bivalves
are often easily recognizable from their irregular
morphology, developed because of the requirement
to conform to substratal irregularities, and are mark-
edly inequivalve. As they tended to evolve from
pleurothetic byssate stock, most are also monomyar-
ian (Figure 7B). Most cement mainly by the right
valve, but a major group, the oysters, do so by the
left valve. The size of the attachment scar varies and
the substrate may be instantly recognizable; for
example, oysters were often attached to ammonite
shells, even if the scar and substrate are no longer
attached. Most cementers have thick, robust shells
and may be extravagantly ornamented with spines or
flanges (e.g., Spondylus; Figure 7B).
Free Living
A number of different taxa have independently aban-
doned attachment to become free living on softer
sediments. Here, the challenge is not to sink into the
substrate, and this has been solved in two ways. The
first is by adopting a ‘snow-shoe’-type morphology,
i.e., resting on a large surface area, epitomized by the
wafer-thin window pane shell Placuna (Figure 7E).
Alternatively, they may be semi-submerged in the soft
sediment like an iceberg. This strategy is inferred for
the thick-shelled devil’s toenail Gryphaea, common in
Mesozoic clay facies (Figure 7F). These ‘aberrant’
oysters clearly had a cemented phase, marked by a
small attachment scar at the umbo. A few free-living
bivalves (notably several groups of scallops) also
have the ability to swim short distances if they are
threatened. These have smooth hydrodynamic shells
and a well-developed posterior adductor muscle
whose vigorous contraction provides the propulsion.
Boring
A number of groups, once again polyphyletically,
have evolved to excavate ‘burrows’ in hard substrates
by boring. The most successful of these, the mytilid
lithophagids, do so principally by acidic secretions
(which presents its own challenge of not dissolving
its own shell), whilst others, e.g., Pholas, bore at least
partly by physically rasping the substrate with small
projections on the outside of the shell. Some of the
most bizarre borers are the teredinids which excavate
long cylindrical boreholes in wood and have the en-
zymatic capability of digesting the cellulose. These
‘shipworms’ are thought to have been part of the
undoing of the ships of the Spanish Armada.
Members of the boring group as a whole have a
very varied morphology, but may be easily recognized
because they are almost invariably fossilized within
Statesman of London, called it "difficult, though not absolutely so." The Times of
London, did an informal poll, asking seven reporters and a math student to read it
and
report on its accessibility. The verdict was mixed. "It all made beautiful sense as
I read it,
though it tended to vanish like a dream when I put the book down," one wrote.
Albert Einstein once said that scientific theories should be able to be described
so simply
that a child could understand them. Complaints that modern physics fails this
standard
abysmally are as old, well, as modern physics, and are not confined to the
childlike
public.
The story goes that when the astronomer Arthur Eddington, whose observations of
light
bending during a solar eclipse in 1919 confirmed Einstein's general theory of
relativity,
was congratulated by a colleague on being one of the three people in the world who
understood the abstruse theory, Eddington fell uncharacteristically silent. Chided
for
exhibiting a false modesty, Eddington replied, on the contrary, that he had been
trying to
imagine who the third person could be.
This newspaper's early accounts of Einstein's and Eddington's 1919 breakthroughs
focused on the theory's incomprehensibility. "Efforts made to put in words
intelligible to
the nonscientific public the Einstein theory of light proved by the eclipse
expedition so
far have not been very successful," began a article on Nov. 10, 1919.
Niels Bohr, one of the founders of quantum mechanics, once said that anyone who was
not outraged on hearing about the theory -- with its waves acting as particles,
particles
acting like waves, and the microscopic randomness and uncertainty it ascribed to
nature -
- had not really understood it.
Recent advances have made it even harder to explain the universe. The latest
version of
the putative theory of everything posits a universe with 10 or 11 dimensions,
instead of
35the 3 of space and 1 of time of everyday experience, inhabited by wriggling
strings or
membranes. Nevertheless, scientists go on gamely trying to tell us what they are up
to, in
a book-writing tradition that includes Darwin's "Origin of Species," and
Einstein's,
"Relativity: The Special and the General Theory," written in 1916 and never out of
print.
Part of the lure of these books is the chance to reclaim one's citizenship in a
troubled and
baffling cosmos by hearing the word from the horse's mouth, from someone who has
touched the cosmic mystery personally. But another part is surely being treated
like an
adult, of entering a rough-hewn colleagueship by being trusted to put work into
deciphering statements like the one at the beginning of this essay, or to deal with
straight
talk of the nature of science and the universe.
Here, for example, is Dr. Hawking about those troublesome extra dimensions required
by
string theory but apparently unavailable for parking cars. "I must say that
personally, I
have been reluctant to believe in extra dimensions," he writes on Page 54 of the
new
book. "But as I am a positivist, the question 'Do extra dimensions really exist?'
has no
meaning. All one can ask is whether mathematical models with extra dimensions
provide
a good description of the universe."
In other words, if the experiments come out right, it doesn't matter. This could be
considered jarring if you cling to the notion that science is the search for a
reality that is
deeper than the measurements on a laboratory table. But, quantum theory and
relativity
have taught us, science is about what can be observed and measured or it is about
nothing
at all. In science, as in democracy, there is no hidden secret knowledge, all that
counts is
on the table, observable and falsifiable. All else is metaphysics.
When it comes to putting the goods on the table without condescending, Dr. Hawking
is a
genius. While many authors of science books plough through chapters full of
fundamentals before getting to the new stuff, Dr. Hawking, with perhaps a
heightened
appreciation of time, breezes speedily to the frontier without apologies.
For those who cannot keep up, Dr. Hawking has also provided a legacy. The success
of
his earlier book and that of Carl Sagan's "Cosmos" are widely credited with having
given
a commercial lift to the science-book genre, helping pave the way for efforts like
"The
Elegant Universe," by Dr. Brian Greene, a Columbia University string theorist; "The
Inflationary Universe," by Dr. Alan Guth, cosmologist at the Massachusetts
Institute of
Technology; and "The Quark and the Jaguar," by the Nobel laureate Murray Gell-Mann.
To the extent that Dr. Hawking's earlier success has spawned imitators and widened
the
circle of readers and their sophistication, he has engineered a kind of positive
feedback,
and he has increased the odds that the readers will follow him and get to the end
of the
book this time.
ENDLESS POSSIBILITIES
The End of Everything
By DENNIS OVERBYE
In the decades that astronomers have debated the fate of the expanding universe --
whether it will all end one day in a big crunch, or whether the galaxies will sail
apart
forever -- aficionados of eternal expansion have always been braced by its
seemingly
endless possibilities for development and evolution. As the Yale cosmologist Dr.
Beatrice
Tinsley once wrote, "I think I am tied to the idea of expanding forever."
Life and intelligence could sustain themselves indefinitely in such a universe,
even as the
stars winked out and the galaxies were all swallowed by black holes, Dr. Freeman
Dyson,
a physicist at the Institute for Advanced Study, argued in a landmark paper in
1979. "If
my view of the future is correct," he wrote, "it means that the world of physics
and
astronomy is also inexhaustible; no matter how far we go into the future, there
will
always be new things happening, new information coming in, new worlds to explore, a
constantly expanding domain of life, consciousness, and memory."
Now, however, even Dr. Dyson admits that all bets are off. If recent astronomical
Transcription Control in
Eukaryotes
Transcription in eukaryotes differs from that in
prokaryotes in two main respects. In eukary-
otes, one gene codes for a single polypeptide
(monocistronic transcription unit) and the ini-
tial transcript is processed into mature mes-
senger mRNA. This involves intron splicing (see
p. 50) and substantial modification of the ends
of the primary transcript.
A. Prototype of a eukaryotic structural
gene
A structural gene is a gene that codes for a poly-
peptide gene product. It can be divided into sec-
tions involved in transcription (transcription
unit) and regulatory sequences. Regulatory
sequences are located both upstream (the 5!
direction) and downstream (the 3! direction) of
the gene. In addition, internal regulatory
sequences may occur in introns. Some regula-
tory sequences are located far from the gene.
Together with the promoter (see p. 206), they
are required to regulate transcription.
nosine is methylated in position 7, as are the
two initial ribose residues at the beginning of
the RNA chain. Except for the mRNAs tran-
scribed by DNA viruses, eukaryotic mRNA usu-
ally contains a single protein-coding sequence
(monocistronic messenger).
D. Polyadenylation at the 3! end
Eukaryotic termination signals have been less
well recognized than the regulators of gene ac-
tivity at the 5! end. Eukaryotic primary tran-
scripts are split by a specific endonuclease
shortly after the sequence AAAUAA. Sub-
sequently, about 100 – 250 adenine nucleotides
are attached to the 3! end of the transcript by
means of a poly(A)-polymerase (polyadenyla-
tion). The poly(A) end binds to a protein. All
mRNAs, except those that code for histone pro-
teins, possess a poly(A) terminus.
(Figures after Singer and Berg, 1991).
References
Singer, M., Berg, P.: Genes & Genomes. Blackwell
Scientific, Oxford, 1991.
B. Prototype of mature eukaryotic
mRNA
Mature eukaryotic mRNA is produced from its
precursor RNA by the removal of introns, addi-
tion of a 5! cap at the 5! end, and addition of
numerous adenine nucleotides at the 3! end
(polyadenylation). A noncoding sequence (5!
leader) is located in front of the translation start
signal (AUG), and a trailer sequence, at the 3!
end in back of the translation stop signal (UAA).
Both addition of the 5! cap and polyadenylation
involve enzymatic reactions.
C. 7-Methyl-guanosine cap
The translation of eukaryotic mRNA is similar to
that of prokaryotic mRNA, with two distinct
differences: (1) transcription and translation
occur at different locations in the eukaryotic
cell: transcription occurs in the cell nucleus,
and translation in the cytoplasm; (2) the 5! and
3! ends of eukaryotic mRNA have special struc-
tures. The structure at the 5! end is called a cap.
Through the action of guanosine-7-methyl-
transferase, guanosine is bound by a tri-
phosphate bridge to the first and second ribose
groups of the precursor mRNA chain. The gua-
Passarge, Color Atlas of Genetics © 2001 Thieme
All rights reserved. Usage subject to terms and conditions of license.Transcription
Control in Eukaryotes
All rights reserved. Usage subject to terms and conditions of license.
215216
Fundamentals
Regulation of Gene Expression
in Eukaryotes
Precisely regulated gene expression is a pre-
requisite for producing and maintaining the
many different types of cells and tissues of a
multicellular organism. Cells differentiate into
their particular cell types by means of combina-
tions of expressed and repressed genes. During
differentiation the tightly regulated genes func-
tion in the order, usually sequential, required
for a particular cell fate (developmental path-
ways). Many regulator genes and their proteins
have been identified (cf. part III, Genetics and
Medicine). The following outlines some impor-
tant principles of the specific control of gene ex-
pression in eukaryotic cells.
A. Levels of control of eukaryotic gene
expression
In principle, expression can be regulated at four
distinct levels. The first and by far the most im-
portant is primary control of transcription.
Processing to mature RNA can be regulated at
the level of the primary RNA transcript. A
frequently observed process is alternative splic-
ing (see D). Translation can be varied by RNA
editing (see B for an example). At the protein
level, posttranslational modifications can de-
termine the activity of a protein. The cleavage of
preproinsulin to form mature insulin, glycosy-
lation or hydroxylation, and protein folding are
some examples (see p. 32, 362).
B. RNA editing
RNA editing modifies genetic information at the
RNA level. An important example is the apolipo-
protein-B gene involved in lipid metabolism. It
encodes a protein of 4538 amino acids, apolipo-
protein B. This is synthesized in the liver and
secreted into the blood, where it transports
lipids. A related shorter form of the protein with
2153 amino acids, Apo B-48 (250 kDa, instead of
512 kDa for Apo B-100), is synthesized in the in-
testine. An intestinal deaminase converts a cy-
tosine in codon 2158, CAA (glutamine), to uracil
(UAA). This change results in a stop codon (UAA)
and thereby terminates translation at this site.
C. Long-range gene activation by an
enhancer
Enhancers control gene activity at a distance.
An enhancer is a distant site involved in initia-
tion of transcription (see p. 206). It may be lo-
cated either upstream or downstream of the
same DNA strand (cis-acting) or on a different
DNA strand (trans-acting). Enhancer elements
provide tissue-specific or time-dependent reg-
ulation. It is unclear how enhancers can exert
their effect from a considerable distance. One
model suggests that DNA forms a loop between
enhancer and promoter. Activator proteins
bound to the enhancer, e.g., a steroid hormone,
could then come into contact with the general
transcription factor complex at the promoter.
Others might function as repressors (cf. tran-
scription control in prokaryotes, p. 210).
D. Alternative RNA Splicing
A DNA segment can code for different forms of
mRNA when different introns are removed from
the primary transcript (alternative splicing). By
means of alternative gene splicing, a gene can
code for different, albeit similar gene products.
This allows a high degree of functional flexi-
bility. Numerous examples of differential RNA
splicing are known for mammalian genes. For
example, the primary transcript for the calci-
tonin gene contains six exons. They are spliced
into two different types of mature mRNA. One,
consisting of exons 1 – 4 (but not exons 5 and 6),
is produced in the thyroid and codes for calci-
tonin. The other consists of exons 1, 2, 3, 5, and
6, but not exon 4. It codes for a calcitonin-like
protein in the hypothalamus (calcitonin gene-
related product, CGRP).
References
Alberts, B., et al: Essential Cell Biology. An Intro-
duction to the Molecular Biology of the Cell.
Garland Publishing Co., New York, 1998.
Lewin, B.: Genes VII, Oxford Univ. Press, Oxford,
2000.
Blackwood E.M., Kadonga J.F.: Going the dis-
tance: A current view of enhancer action.
Science 281 :60 – 63, 1998.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freeman &
Co, New York, 1995.
Watson J.D., et al.: Molecular Biology of the
Gene. 4 th ed. Benjamin/Cummings Publish-
ing Co., Menlo Park, California, 1987.
All rights reserved. Usage subject to terms and conditions of license.217
Regulation of Gene Expression in Eukaryotes
1
Apo B-100
Glu
CAA
Cytosol
5'
mRNA unedited
Nucleus
DNA
UAA
5'
3'
Cytosine deamination
by intestinal deaminase
3'
2158
Apo B-84
B. RNA editing
Activator protein
Control of
processing
(alternative
splicing)
Transcription
start site
5'
3'
Enhancer
mRNA
Promoter
Binding of an
activator protein
to the transcription
complex
Control of
translation
(mRNA editing)
Enhancer
Protein
Activator
protein
Transcription factors
Control of
protein activity
active
Translation
stop
1
Control of
transcription
Primary transcript
4538
inactive
A. Levels of control of eukaryotic
gene expression
Transcription
Promoter with
transcription
factors and
RNA polymerase II
C. Long-range gene activation by an enhancer
Calcitonin gene
5'
Exon 1
Exon 2
Exon 3
Primary RNA transcript
5'
1
mRNA
5'
2
3
C cells in thyroid
1
2
Exon 4 Exon 5
4 5
Exon 6 3'
6 3'
Transcription
RNA processing
3
4
3'
5'
Translation
Calcitonin
Hypothalamus
1
2
3
5
6
Translation
Different
gene products
D. Alternative RNA splicing
CGRP
(Calcitonin
gene-related
peptide)
3'218
Fundamentals
DNA-Binding Proteins
Regulatory DNA sequences interact with pro-
teins to exert proper functional control. Regula-
tory proteins can recognize specific DNA
sequences because the surface of the proteins
fits precisely onto the DNA surface. Three basic
groups of regulatory DNA sequences can be dis-
tinguished: (1) sequences that establish the
exact beginning of translation; (2) DNA seg-
ments that regulate the end, or termination;
and (3) DNA sequences near the promoter that
have specific effects on gene activity (repres-
sors, activators, enhancers, and others).
A. Binding of a regulatory protein to
DNA
Gene regulatory proteins can recognize DNA
sequence information without having to open
the hydrogen bonds within the helix. Each base
pair represents a distinctive pattern of hydro-
gen bond donors (example shown in red) and
hydrogen acceptors (example shown in green).
These proteins recognize the major groove of
DNA, where binding takes place. Here a single
contact of an asparagine (Asn) of a gene-regula-
tory protein with a DNA base adenine (A) is
shown. A typical area of surface-to-surface con-
tact involves 10 – 20 such interactions. (Figure
redrawn from Alberts et al., 1998, p. 276).
B. An α helix inserts into a major
groove of operator DNA
One part of the protein, an α helix (the
sequence-reading or recognition helix) is in-
serted into the major groove of DNA. Here the
sequence Q-Q-Q-S-T (glutamine Q, serine S,
threonine T) in the recognition sequence of the
bacteriophage 434 repressor bonds with
specific bases in a major groove of operator
DNA. (Figure redrawn from Lodish et al., 2000,
p. 351).
C. Zinc finger motif
Another group of proteins are called zinc fingers
because they resemble fingers (see D). They are
involved in important functions during embry-
onic development and differentiation. The basic
zinc finger motif consists of a zinc atom con-
nected to four amino acids of a polypeptide
chain. Here, two histidine (H) and two cysteine
(C) residues are shown in the schema on the
left. The three-dimensional structure on the
right consists of an antiparallel β sheet (amino
acids 1 – 10), an α helix (amino acids 12 – 24),
and the zinc connection. Four amino acids, cys-
teines 3 and 6 and histidines 19 and 23, are
bonded to the zinc atom and hold the carboxy
(COOH) end of the α helix to one end of the β
sheet. (Figure redrawn from Alberts et al., 1994,
p. 411).
D. Zinc finger proteins bind to DNA
The interaction with DNA is strong and specific.
Each protein recognizes a specific DNA
sequence. As the number of zinc fingers can be
varied, this type of DNA-binding has great evo-
lutionary flexibility. (Figure redrawn from Al-
berts et al. 1994).
E. Binding to a response element
Many hormones and growth factors activate
cell-surface receptors. In contrast, steroid hor-
mones enter the target cells and interact there
with a specific receptor protein in the cytosol.
The hormone–receptor complex then migrates
to specific sites of DNA. The hormone-binding
domain will prevent binding to DNA unless the
hormone is present. Activated receptors bind to
specific DNA sequences called hormone re-
sponse elements (HREs). Each polypeptide
chain of the receptor contains a zinc atom
bound to four cysteines (1). The skeletal model
shows the two DNA-binding domains binding
to the HRE in two adjacent major grooves of the
target DNA (2). The space-filling model shows
how tightly the recognition helix of each dimer
of this protein fits into the major groove of DNA
shown in red and green (3). (Figure redrawn
from Stryer, 1995, p. 1002).
References
Alberts, B., et al.: Molecular Biology of the Cell.
3 rd ed. Garland Publishing Co., New York,
1994.
Alberts, B., et al.: Essential Cell Biology. Garland
Publishing Co., New York, 1998.
Lodish, H., et al.: Molecular Cell Biology. 4 th ed.
Scientific American Books, F.H. Freeman &
Co., New York, 2000.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freeman &
Tjian, R.: Molecular machines that control
genes. Sci. Am. 272 :38 – 45, 1995.
All rights reserved. Usage subject to terms and conditions of license.DNA-Binding
Proteins
219
DNA-binding protein
Asn
Major groove
Donor
CH 3
T
H
CH 2
C
H
O N
N H
N
N
N
A
To sugar
25
N
R
K
V
H
Q
N
S
T
To sugar
H
Minor groove
A. Binding of a regulatory protein to DNA
HOOC
Q
H
N
O
Q
H Acceptor
H
HN
O
23
3
C
Zn
Q
K
1
Y
B. An ! helix inserts into a major
groove of operator DNA
25
NH 2
HOOC
His
23
C
L
6 C
H 19
R
E
S
L R
A S
His 19
F 10
S
K
V
Zn
E
12
12
D. A zinc finger protein binds to DNA
Cys
443
Cys
440
Zn
1.
Cys
457
2.
Cys
3
1
H 2 N
Zn
C. Zinc finger motif
Cys
460
Cys
6
Zn
3.
E. Binding to a response element
10220
Fundamentals
Other Transcription Activators
Transcription activators are dimeric proteins
with distinct functional domains: a DNA-bind-
ing domain and an activation domain. The DNA-
binding domain interacts with specific regula-
tory DNA sequences. The activation domain in-
teracts with other proteins that stimulate tran-
scription. Transcription activators participate in
the assembly of the initiation complex, for ex-
ample, by stimulating the binding of transcrip-
tion factor IID (TFIID, see p. 212) to the pro-
moter. Other activators may interact with
general transcription factors. They provide a
second level of transcriptional control.
A. Leucine zipper dimer
Most DNA-binding regulatory proteins recog-
nize specific sites as dimers. One part of the
molecule serves as the recognition molecule,
the other stabilizes the structure. A particularly
striking example is given by proteins with a
leucine zipper motif. The name is derived from
the basic structure. Two α helices are joined like
a zipper by periodically repeated leucine resi-
dues located at the interface of the two helices.
The two helices separate, form a Y-shaped
structure, and extend into the major groove of
the DNA (1). Leucine zipper proteins may be ho-
modimers with identical subunits (2, 3) or het-
erodimers with different albeit similar subunits
(4). The ability to form unlike dimers (hetero-
dimerization) greatly expands the spectrum of
specificites. The use of combinations of differ-
ent proteins to control cellular functions is
called combinatorial control. (Figure redrawn
from Alberts et al., 1994).
A DNA-binding motif related to the leucine zip-
per is the helix–loop–helix (HLH) motif (not
shown). The HLH motif consists of one short α
helix and one longer α helix. The two α helices
are connected by a flexible loop of protein.
ment are regulated by steroids (steroid-respon-
sive transcription). The latter include glucocor-
ticoids and mineralocorticoids, the steroids of
glycogen and mineral metabolism; sex hor-
mones, which function in embryonic sex differ-
entiation and control of reproduction; and
others. Normal bone development and function
are under the control of steroidlike vitamin D.
Another steroidlike hormone is retinoic acid, an
important regulator of differentiation during
embryogenesis (morphogen). These hormones
initiate their physiological effects by associa-
tion with corresponding steroid-specific trans-
cellular receptors (hormone–receptor com-
plex).
C. Evidence of a protein-binding
region in DNA
Protein-binding regions in DNA represent regu-
latory areas; thus, their analysis can yield some
insights into gene regulation. Protein-binding
DNA regions can be demonstrated in several
ways. With band-shift analysis (1), protein-
bound and non-protein-bound DNA fragments
are differentiated using gel electrophoresis in
direction towards the small fragments, a DNA
fragment that is part of a DNA-protein complex
migrates more slowly than a free DNA fragment
of the same size. The DNA-protein complex is
found at a different position (“band shift”). DNA
footprinting (2) is another procedure for identi-
fying protein-binding sites on DNA. The prin-
ciple of DNA footprinting is that a protein-
bound DNA region, e.g., the polymerase-pro-
moter complex, is protected from the effects of
a DNA-cleaving enzyme (DNAase I). Previously
isolated DNA is cut into different fragments by
DNAase, and the fragments are sorted accord-
ing to size by gel electrophoresis. Since the DNA
protein-binding region is protected from cleav-
age by DNAase I (DNAase I protection experi-
ment), DNA bands from the binding region are
missing (“footprint”).
B. Activation by steroid hormone
binding References
Transcriptional enhancers are regulatory re-
gions of DNA that increase the rate of transcrip-
tion. Their spacing and orientation vary relative
to the starting point of transcription. An en-
hancer is activated by binding to a hormone–re-
ceptor complex. This activates the promoter,
and transcription begins (active gene). Numer-
ous important genes in mammalian develop- Alberts, B., et al.: Molecular Biology of
the Cell.
1994.
Alberts, B., et al.: Essential Cell Biology. Garland
Publishing, Co., New York, 1998.
Lodish, H., et al.: Molecular Cell Biology. 4 th ed.
Scientific American Books, F.H. Freeman &
All rights reserved. Usage subject to terms and conditions of license.Other
Transcription Activators
221222
Fundamentals
Inhibitors of Transcription and
Translation
A number of natural and artificial substances
are able to inhibit transcription or translation.
They can be used to treat cancer or as antibiotics
to treat infections. Although most substances
are unspecific and not suitable for therapeutic
purposes, some are very specific and therefore
important for the understanding of transcrip-
tion and translation or for therapy. Basically,
one can distinguish whether an agent interferes
with transcription or with translation.
A. Insertion of actinomycin D
between a GC base pair
Actinomycin D is a complex polypeptide pro-
duced by a species of streptomyces bacteria. It
consists of a phenoxazone ring with two sym-
metrical side chains (1). It acts by becoming in-
tercalated between two neighboring GC base
pairs in double-stranded DNA. Viewed from the
side (2), the inserted actinomycin D molecule is
seen very distinctly within the DNA double
helix. In the view from above (3) the actinomy-
cin D molecule forms a narrow layer within the
DNA double helix, bound by the two neighbor-
ing GC base pairs. The degree of inhibition by
actinomycin D varies greatly. High concentra-
tions of actinomycin D block replication,
whereas low concentrations suffice to inhibit
transcription.
bosome, and the protein synthesis ends prema-
turely.
(Figures A and B from Singer and Berg, 1991).
C. Inhibitors of protein synthesis
Numerous naturally occurring and artificially
produced substances inhibit protein synthesis
by inhibiting transcription or certain phases of
translation. Some have clinical significance as
antibiotics; others are toxicologically signifi-
cant. An example for the specificity of some in-
hibitors is α -amanitin, a dicyclic octapeptide of
the fungus Amanita phalloides. In very low con-
centrations, it binds to RNA polymerase II and
thereby blocks the formation of precursor
mRNA in eukaryotes. In contrast, RNA poly-
merase I is insensitive to this toxin, and poly-
merase III binds to it only in higher concentra-
tions. (Data after Singer and Berg, 1991).
References
Singer, M., Berg, P.: Genes & Genomes. Blackwell
Scientific, Oxford, 1991.
B. Puromycin imitates an aminoacyl
tRNA
Puromycin, a polypeptide from Streptomyces al-
boniger, blocks polypeptide synthesis in the ri-
bosomes of prokaryotes and eukaryotes. Its ac-
tion is based on the structural similarity with an
aminoacyl tRNA. An aminoacyl tRNA is a tRNA
molecule with an amino acid attached to its 3!
end. Normally a peptide bond is formed by pep-
tidyltransferase between the amino group of
the incoming aminoacyl tRNA at the A (amino-
acyl) position and the carboxyl group of the
peptidyl tRNA at the P (peptidyl) position. The
structure of puromycin resembles that of
aminoacyl tRNA, but lacking an interaction with
the codon it cannot be attached to the A posi-
tion in the ribosome. The resulting polypep-
tidyl–puromycin adduct is removed from the ri-
All rights reserved. Usage subject to terms and conditions of license.Inhibitors of
Transcription and Translation
223224
Fundamentals
DNA Methylation
Methylation of cytosine residues in DNA plays
an important role in gene regulation. DNA
methylation is required for normal embryonic
development. Genomic imprinting, X chromo-
some inactivation, chromatin modification, and
silencing of endogenous retroviruses all depend
on establishing and maintaining proper meth-
ylation patterns. DNA methylation is gene-
specific and occurs genome-wide. Two types of
methyltransferase can be distinguished by their
basic functions: maintenance methylation and
de novo methylation.
A. Maintenance methylation
This type of methylation is responsible for
adding methyl groups to the newly synthesized
DNA strand after replication and cell division.
The methylated sites in the parental DNA (1)
after replication (2) serve as template for cor-
rect methylation of the two new strands. This
ensures that the previous methylation pattern
is correctly maintained (3). It results in both
daughter strands being methylated at the same
sites as the parental DNA. The enzyme re-
sponsible for this is Dnmt1 (DNA methylase 1).
Its function is essential. Mice deficient in this
enzyme die as a result of genome-wide de-
methylation.
B. Recognition of a methylated DNA
segment
Certain restriction enzymes do not cleave DNA
when their recognition sequence is methylated
(1). The enzyme HpaII cleaves DNA only when
its recognition sequence 5!-CCGG-3! is not
methylated (2). MspI recognizes the same 5!-
CCGG-3! sequence irrespective of methylation
and cleaves DNA at this site every time. This
difference in cleavage pattern results in DNA
fragments of different sizes serves to distin-
guish the methylation pattern of the DNA.
C. DNA methylation de novo
This is the second type of DNA methylation.
Here methyl groups are added at new positions
of both strands of DNA, not just in the hemi-
methylated strand as in maintenance methyla-
tion shown in A. Two genes for different
methyltransferases with overlapping functions
in global remethylation have been identified re-
cently: Dnmt3a and Dnmt3b. Unmethylated
DNA (1) is methylated by their enzymes (2) in a
site-specific and tissue-specific manner (3).
Targeted homozygous disruption of the mouse
Dnmt3a and Dnmt3b genes results in severe
developmental defects. Double homozygous
mutants die before day 11.5 of the 21-day
embryonic development.
D. Human DNMT3B gene
Mutations in the human gene DNMT3B encod-
ing type 3B de novo methyltransferase cause a
distinctive disease called ICF syndrome (im-
munodeficiency, centromeric chromosomal in-
stability, and facial anomalies, McKusick catalog
no. 242860). The centromeres of chromosomes
1, 9 and 16, where satellite DNA types 2 and 3
are located, are unstable. Classical satellite DNA
is grossly undermethylated in all tissues. The
human gene (1) consists of 23 exons spanning
47 kb of genomic DNA. Six exons are subject to
alternative splicing. The protein (2) consists of
845 amino acids with five DNA methyl-
transferase motifs (I, IV, V, IX, X) in the C-termi-
nal region. The arrows point to six different mu-
tations. The mutation at position 809 (3), a
change of A to G in codon 809, i.e., GAC (Asp) to
GGC (Gly), leads to the replacement of as-
paragine (Asn) by glycine (Gly). Both parents
are heterozygous for this mutation.
(Figure adapted form Xu et al., 1999).
References
Bird, A: DNA methylation de novo. Science
286 :2287 – 2288, 1999.
Hansen, R. S. , et al.: DNMT3B DNA methyl-
transferase gene is mutated in the ICF im-
munodeficiency syndrome. Proc. Nat. Acad.
Sci. 96 :14 412 – 14 417, 1999.
Okano, M., et al.: DNA Methyltransferases
Dnmt3a and Dnmt3b are essential for de
novo methylation and mammalian develop-
ment. Cell 99 :247 – 257, 1999.
Reik, W., Kelsey, G., Walter, J.: Dissecting de novo
methylation. Nature Genet. 23 : 380 – 382,
1999.
Robertson, K.D., Wolffe, A.P.: DNA methylation
in health and disease. Nature Reviews 1 :11 –
19, 2000.
Xu, G., Bestor, T., et al.: Chromosome instability
and immunodeficiency syndrome caused
by mutations in a DNA methyltransferase
gene. Nature 402 :187 – 191, 1999.
All rights reserved. Usage subject to terms and conditions of license.DNA
Methylation
225226
Fundamentals
Genomic Imprinting
Genetic contributions from both the maternal
and the paternal sets of chromosomes are ne-
cessary for a mammalian zygote to develop nor-
mally. The reason lies in a parent-of-origin-
specific expression of certain genes. The
genome contains defined regions where only
the maternal gene copy is expressed, and not
the paternal copy, and vice versa. This allele-
specific gene expression, depending on the
parental origin, results from so-called genomic
imprinting. Genomic imprinting has important
implications for human genetic disease (see
p. 398).
A. The importance of two different
parental genomes
Different developmental results are observed
when the female pronucleus is removed from a
mouse zygote (1) before the pronuclei have
fused and is replaced either by another male
pronucleus (2) or by a control (i.e., the same as
removed) (3), or when the male pronucleus is
removed and replaced by a female pronucleus
(4) or a control. If the female pronucleus is re-
placed by a male pronucleus (2), the zygote first
appears normal. However, after implantation
should ensue, nearly all androgenotes fail to
complete preimplantation (2). Those few that,
rarely,
reach
postimplantation
develop
completely abnormally. Such embryos do not
develop beyond the 12-somite stage.
When a male pronucleus is replaced by a female
pronucleus (4), a gynogenetic zygote, which
differs markedly from the androgenote. Here,
about 85% of gynogenotes reach normal preim-
plantation development. But although the em-
bryo at first develops fairly well, the extra-
embryonic membranes are absent or under-
developed, and the gynogenote dies before
reaching the 40-somite stage. (Figure redrawn
from Sapienza and Hall, 1995).
B. Human embryonic development
depends on the presence of a
maternal and a paternal genome
A naturally occurring human androgenetic zy-
gote is a hydatidiform mole (1). This is an abnor-
mal placental formation containing two sets of
paternal chromosomes and none from the
mother. No embryo develops although implan-
tation takes place. The placental tissues develop
many cysts (2). When only maternal chromo-
somes are present, an ovarian teratoma with
many different types of fetal tissues develops
(3). No placental tissue is present in this nat-
urally occurring gynogenetic zygote. In tri-
ploidy, a relatively frequent global chromo-
somal lethal human disorder (see p. 402), ex-
treme hypoplasia of the placenta and fetus is
observed when the additional chromosomal set
is of maternal origin (4). (Photographs kindly
provided by Professor Helga Rehder, Marburg.)
C. Genomic imprinting is established in
early embryonic development
The imprint pattern present in somatic cells (1),
with one allele active only either, the maternal
or the paternal, propagated through all mitotic
divisions. However, in primordial germ cells the
imprint is erased (2). During gametogenesis the
imprint is reset (3). In the male germline all
gametes receive the paternal imprint and in the
female germline all gametes receive the mater-
nal imprint. After fertilization, the correct im-
print pattern is present in the zygote (4). It is
maintained through all subsequent cell divi-
sions under the control of a regional imprinting
center (see p. 398).
References
Barlow, D.P.: Gametic imprinting in mammals,
Science 270 :1610 – 1613, 1995.
Horsthemke, B.: Genomisches Imprinting beim
Menschen: Grundlagen und klinische Rele-
vanz. Biospektrum 4 :23 – 26, 1998.
Morrison, I.M., Reeve, A.E.: Catalogue of im-
printed genes and parent-of-origin effects
in humans and animals. Hum. Mol. Genet.
7 :1599 – 1609, 1998.
Reik, W., Surani, A., eds.: Genomic Imprinting.
IRL Press at Oxford University Press, Oxford,
1997.
Sapienza, C., Hall, J.G.: Genetic imprinting in
human disease. pp. 437 – 458. In: The Meta-
bolic and Molecular Bases of Inherited Dis-
ease. 7 th ed. C.R. Scriver, et al., eds. McGraw-
Hill, New York, 1995.
Surani, A.: Imprinting and the initiation of gene
silencing. Cell 93 :309 – 312, 1998.
Tilghman, S. M.: The sins of the fathers and
mothers: Genomic imprinting in mam-
malian development. Cell 96 :185 – 193,
1999.
munication between cells to assure growth,
differentiation, specific functions in different
types of cells, and proper response to external
stimuli. Specific cell–cell interactions between
different types of cells have evolved. A common
leitmotif is the specific binding of an extracellu-
lar signaling molecule (ligand) to a specific re-
ceptor of the target cell to trigger a specific
functional response. The vast variety of
molecules involved in the many different types
of cells can be classified into families of related
structure and function (see Lodish et al., 2000;
Alberts et al., 1994). Two areas are selected
here: the main intracellular functions control-
ling growth and the receptor tyrosine kinases.
A. Main intracellular functions
controlling growth
Growth factors are a large group of different ex-
tracellular molecules that bind with high speci-
ficity to cell surface receptors (1). Their binding
to the receptor (2) activates intracellular signal
transduction proteins (3). This initiates a cas-
cade of events resulting in activation of other
proteins (often by phosphorylation) that act as
second messengers (4). Hormones of different
types are a heterogeneous class of signaling
molecules (5). They enter the cell either by dif-
fusion through the plasma membrane or by
binding to a cell surface receptor (6). Some hor-
mones require an intranuclear receptor (7).
Eventually the signal cascade results in activa-
tion or inactivation of transcription factors (8).
Before transcription and translation ensue, an
elaborate system of DNA damage recognition
and repair systems (9) make sure that cell pro-
liferation is safe (cell cycle control, 10). In the
event that faults in DNA structure have not been
repaired prior to replication, an important
pathway sacrifices the cell by apoptosis (cell
death, 11). (Figure adapted from Lodish et al.,
2000.)
B. Receptor tyrosine kinase family
Like the G protein-coupled receptors (GPCRs,
see p. 268) and their effectors, the receptor ty-
rosine kinases (RTKs) are a major class of cell
surface receptors. Their ligands are soluble or
membrane-bound growth factor proteins. RTK
signaling pathways involve a wide variety of
other functions. Mutations in RTKs may send a
proliferative signal even in the absence of a
growth factor, resulting in errors in embryonic
development and differentation (congenital
malformation) or cancer. Of the more than
twenty different RTK families, five examples are
selected here: the epidermal growth factor re-
ceptor (EGFR); insulin receptor (IR); fibroblast
growth factor receptor (FGFR) types 1, 2, and 3;
platelet-derived growth factor (PDGFR); and
RET (rearranged during transformation).
These receptors share structural features, al-
though they differ in function. All have a single
transmembrane domain and an intracellular ty-
rosine kinase domain of slightly varied size. The
extracellular domains consist of evolutionarily
conserved motifs: cystein-rich regions, im-
munoglobulin (Ig)-like domains, fibronectin re-
peats in the tyrosine kinase with Ig and the EGF.
RTK mutations cause a group of important
human diseases and malformation syndromes.
The phenotypes of the mutations differ accord-
ing to the particular type of RTK involved and
the type of mutation.
References
Alberts, B., et al.: Molecular Biology of the Cell.
1994.
Cohen, M.M.: Fibroblast growth factor receptor
mutations, pp. 77 – 94, In: M.M. Cohen Jr.,
R.E. MacLean, eds., Craniosynostosis, Diag-
nosis, Evaluation, and Management. 2 nd ed.
Oxford University Press, Oxford, 2000.
Lodish, H., et al.: Molecular Cell Biology (with an
animated CD-ROM). 4 th ed. W.H. Freeman &
Muenke, M., et al.: Fibroblast growth factor re-
ceptor–related skeletal disorders: cranio-
synostosis and dwarfism syndromes, pp.
1029 – 1038, In: J.L. Jameson, ed., Principles
of Molecular Medicine. Humana Press, To-
towa, New Jersey, 1998.
Münke, M., Schell, U.: Fibroblast-growth-factor
receptor mutations in human skeletal dis-
orders. Trends Genet. 11 : 308 – 313, 1995.
Roberton, S. C., Tynan, J.A., Donoghue, D.J.: RTK
mutations and human syndromes: when
good receptors turn bad. Trends Genet.
16 : 265 – 271, 2000.
All rights reserved. Usage subject to terms and conditions of license.Intracellular
Signal Transduction Systems
265266
Genetics and Medicine
Types of Cell Surface Receptors
Specific receptors on cell surfaces (and in the
nucleus or cytosol) convey cell-to-cell signals
into the cells and the functional answers. The
basic structures of their genes are similar be-
cause they have been derived from a relatively
small group of ancestral genes. They way they
bind to the ligand (the signal-releasing
molecule) and the functional answer of the cell
are specific. When a ligand binds to a receptor, a
series of reactions is initiated that alters the
function of the cell. Receptors with direct and
indirect ligand effects can be distinguished. Epinephrine, norepinephrine, and
histamine
act directly and very rapidly. Peptide hormones
such as insulin or adrenocorticotropic hormone
(ACTH) initially occur as precursor polypep-
tides, which are split by specific proteases to
form active molecules. Some peptide hormones
are coded for by a common gene; differential
RNA splicing of the transcript of this gene pro-
duces different precursors for translation.
(Abbreviations used: ACTH, adrenocorti-
cotropic hormone; FSH, follicle-stimulating
hormone; LH, leutinizing hormone; TSH, thy-
roid-stimulating hormone.) (Figure data after
Lodish et al., 2000.)
A. Cell surface receptors with direct
ligand effect C. Cell surface receptors with indirect
Therefore, if a magnitude equal to Z is applied to each segment of LH
in. such a way that its centre of gravity is at the centre of the segment,
Elll the magnitudes 1 will be equal to A. Further, the centre of gravity
of the magnitude made up of all these magnitudes will he the point E,
remembering that they are an even numher and that LE is equal to HE
rrClJ}{H;lLIUn V).
Similarily it could he shown that if a magnitude
to Z was applied to each of the segments of KH, with its centre
.nr.....
at the centre of each segment, all those magnitudes 1 would
to B and that the combined centre of gravity would he D.
~4']I"'_'I"''I'.I'I.a. A is applied at the point E and the magnitude B at
·.In.er~~to]re certain equal magnitudes are placed on the
centres of gravity are separated from each other by the
an even number. It is therefore clear that
magnitude composed of all these magnitudes
..u.:&"'~.""JL'U ~J.L the line on which the centres of gravity of
rrO'DO~;ltl()n V). But the length LE is
EC to the length CK. Thus the
• lQl~lgIlllttlde composed. of aU these areas is the
ma~].tu{le A is applied to the •. point E and
the two areas will be in equilibrium
r:!I"I:rlI· ... "I:T
e~1te.ld.ela..· thiS J)r~(}D~).slltlOn to. the. case
of magnitudes
demonstration depended
reproduced this proof of
I
Read, "the combination of all these magnitudes. "28
THE ORIGINS
Proposition VI in its entirety in order to illustrate the nature of Archi-
medes' logical apparatus. This should not be allowed, however, to
create too great an illusion of power.
Indeed, Archimedes assumes in this proof that the load on the
fulcrum of a lever is equal to the sum of the two weights which it
supports.! Further, he made use of the principle of superposition
of equilibrium states, without emphasising that this was an experi-
mental postulate. Finally, and this is the most telling objection to
the preceeding analysis, Archimedes, together with those of his suc-
cessors who tried to improve his proof, tacitly made the hypothesis
that the product PL measures the effect of a weight P placed at a
distance L from a horizontal axis. In fact, in the case of complete
symmetry which is envisaged in Archimedes' first postulate, equili-
brium obtains whatever law of the form Pf(L)is taken as a measure
of the effect of the weight P. It is therefore impossible, with the help
of Archimedes' postulates alone, to substantiate Proposition VI in a
logical way.2
For the rest, the treatise On the Equilibrium of Planes is concerned
with the determination of the centres of gravity of particular geome-
trical figures. After having obtained the centres of gravity of a tri-
angle, a parallelogram and a trapezium, Archimedes determined the
centre of gravity of a segment of a parabola by means of an analysis
which is a milestone in the history of mathematics (Book II, Pro-
position VIII).
We shall now concern ourselves with Archimedes' treatise on
Floating Bodies. The author starts from the followillg hypothesis-
" The nature of a fluid is such that if its parts are equivalently
placed and continuous with each other, that which is the least compress-
ed is driven along by that which is the more compressed. Each part
;of the fluid is compressed by the fluid which is above it in a vertical
dir~ction, whether the fluid is falling some,vhere or whether it is
driven from one place to another. "
From this starting point, the following propositions derive in a
logical sequence.
Proposition I. - If a surface is intersected by a plane which always
passes through the same point and if the section is a circumference
(of a circle) having this fixed point as its centre., the surface is that
of a sphere.
1 This is a point which can be established rigorously by considerations of symmetry
alone, as FOURIER was to show, much later, in his perfection of a similar attempt
due
to n'ALEMBERT.
2 Cf MACH, Mechanics, p. 21.
Throughout this work, Mach's treatise ,vill be
indicated by the letter ]}f.29
HELLENIC SCIENCE
Proposition II. - The surface of any fluid at rest is spherical and
the centre of this surface is the same as the centre of the Earth.
This result had already, as we have seen, been enunciated by Aristotle.
Proposition III. - If a body whose weight is equal to that of
the same volume of a fluid (a) is immersed in that fluid, it will sink
until no part of it remains above the surface, but will not descend
further.
We shall reproduce the proof of this proposition, which is the
origin of Archimedes' Principle.
" Let a body have the same heaviness as a liquid. If this is possi-
ble, suppose that the body is placed in the fluid with part of it above
the surface. Let the fluid be at rest. Suppose that a plane which
passes through the centre of the Earth intersects the fluid and the
K
D
Fig. 3
body immersed in it in such a ,yay that the section of the fluid is ABeD
and the section of the body is EHTF. Let [( be the centre of the
Earth, BHTC be the part of the body which is immersed in the fluid
and BEFC the part which projects out of it. Construct a pyramid
whose base is a parallelogram in the surface of the fluid (a) and whose
apex is the centre of the Earth. Let the intersection of the faces
of the pyramid by the plane containing the arc ABCD be KL and
[(M. In the fluid, and below EFTH draw another spherical surface
XOP having the point I( as its centre, in such a way that XOP is the
section of the surface by the plane containing the arc ABCD. Take
another pyramid equal to the first, with which it is contiguous and
continuous, and such that the sections of its faces are KM and KN.
Suppose that there is, in the fluid, another solid RSQY which is made
of the fluid and is equal and similar to BHTC, that part of the body30
THE ORIGINS
EHTF which is immersed in the fluid. The portions of the fluid
which are contained by the surface XO in the first pyramid and the
surface OP in the second pyramid are equally placed and continuous
with each other. But they are not equally compressed. For the
portions of the fluid contained in XO are compressed by the hody
EHTF and also by the fluid contained by the surfaces LM, XO and
those of the pyramid. The parts contained in PO are compressed
hy the solid RSQ Y and by the fluid contained by the surfaces OP, MN
and those of the pyramid. But the weight of the fluid contained
between MN and OP is less than the combined heaviness of the fluid
between LM and XO and the solid. For the solid RSQY is smaller
than the solid EHTF-RSQY is equal to BHTC-and it has heen
assumed that the hody immersed has, volume for volume, the same
heaviness as the fluid. If therefore one takes away the parts which
are equal to each other, the remainder will be unequal. Consequently
it is clear that the part of the fluid contained in the surface OP will
be driven along by the part of the fluid contained in the surface XO,
and that the fluid will not remain at rest. Therefore, no part of the
hody which has been immersed will remain above the surface. How-
ever, the body will not fall further. For the body has the same
heaviness as the fluid and the equivalently placed parts of the fluid
compress it similarily."
Proposition IV. - If a body which is lighter than a fluid is placed
in this fluid, a part of the 'body will remain above the surface.
(Proof analogous to that of Proposition III.)
Proposition V. - If a body which is lighter than a fluid is placed
in the fluid, it will he immersed to such an extent that a volume of
:fluid which is equal to the volume of the part of the body immersed
h.as the same weight as the whole body.
The diagram is the same as the preceding one (Proposition III).
" Let the liquid be at rest and the body EHTF be lighter than
the fluid. If the fluid is at rest, parts which are equivalently placed
,vill be similarly compressed. Then the fluid contained by each of
the <_§urfaces XO and OP is compressed by an equal weight. But,
if the body BHTC is excluded, the weight of fluid in the first pyramid
is equal, with the exclusion of the fluid RSQY, to the ,veight of fluid
in the second pyramid. Therefore it is clear that the weight of the
body EHTF is equal to the weight of the fluid RSQY. From which
it follows that a volume of fluid equal to that of the body which is
immersed has the same weight as the whole body."HELLENIC SCIENCE
31
Proposition VI. - If a body which is lighter than a fluid is totally
and forcibly immersed in it, the body will be thrust upwards with
a force equal to the difference between its weight and that of an equal
volume of fluid.
Proposition VII. - If a body is placed in a fluid which is lighter
than itself, it will fall to the bottom. In the fluid the body will be
lighter by an amount which is the weight of the fluid which has the
same volume as the body itself.
The first Book of the treatise On Floating Bodies concludes with
the following hypothesis- (,(, We suppose that bodies which are thrust
upwards all follow the vertical which passes through their centre of
gravity. "
In Book II, Archimedes modified the principle which is the subject
of Proposition V, Book I, to the following form-
" If any solid magnitude which is lighter than a fluid is immersed
in it, the proportion of the weight of the solid to the weight of an
equal volume of fluid will be the same as the proportion of the volume
of that part of the solid which is submerged to the volume of the whole
solid. "
We shall pass over the proof of this proposition, in which Archimedes
once more deploys that powerful logical apparatus with which we are
now familiar. The rest of Book II is devoted to a detailed study of
the equilibrium of a floating body which is shaped like a right segment
of a (,(, parabolic conoid." In Archimedes' language (in the treatise
On Conoids and Spheroids), this term refers to the figure which we
would now call a parabolic cylinder. It may be surmised that Archi-
medes was interested in this problem for a most practical reason, for
this surface is similar to that of the hull of a ship.
It is of interest that, throughout this study, Archimedes approxi-
mated the free surface of a fluid by a plane, and that he treated verticals
as if they were parallel. This is necessary if the concept of centre
of gravity is to be utilised. Thus Archimedes must have understood
the necessity and the practical importance of this approximation,
even though his principle was based on the convergence of the verticals
at the centre of the Earth, the spherical symmetry of fluid surfaces
and a rather vague hypothesis about the pressures obtaining in the
interior of a fluid.CHAPTER TVVO
ALEXANDRIAN SOURCES
1.
~t\ND
ARABIC MANUSCRIPTS
THE " MECHANICS" OF HERO OF ALEXANDRIA.
It seems that Hero of Alexandria lived at some time during the
lInd Century A. D. His treatise Mechanics discusses certain simple
machines-the lever, pulley-block and the screw-alone or in various
combinations, and is only available to us in the form of an arabic version
which has been translated and published by Carra de Vaux. 1
As far as it concerns the history of mechanics, the essential import-
ance of this work lies in the fact that its author used the now classical
idea of moment in his discussion of a lever which was not straight.
Whether or not this conception was an original one remains doubtful.
Indeed alexandrian learning had access to a treatise of Archimedes that
was devoted to levers (lleel evywv) and in which the problem of the
angular lever had been treated. No trace
of this writing is extant.
However this may he, we shall quote
from Hero's Mechanics.
"Consider all arm of a balance which
does not have the same thickness or
heaviness throughout its length. It may
he made of any material. It is in equili-
brium when suspended from the point 'Y
-by equilibrium we understand the arrest
of the beam in a stahle position, even
e
though it may be inclined in one direction
Fig. 4
or the other. Now let weights he suspended
at some points of the beam-sa)'" at ()
and B. The beam will take up a new position of equilibrium after the
weights have heen hung on. Archimedes has shown, in this case as
1
Journal asiatique, 1894.ALEXANDRIAN SOURCES AND ARABIC MANUSCRIPTS
mW]r'se .&.v...
,P,"....
33
"'''''''''''''''.VLL of the weights to each other is the same as the
VLL of the :respective distances. l Concerning these distances
irregular and inclined heams, it should he imagined that
allowed to fall from y towards the point ,. Construct a
passes through the point '-the line 11'O-and ""'"hich should
arranged to intersect the string at right angles. When the beam is
at rest the relation of'1J to CO is the same as the relation of the weight
hung at the point B to the ,veight hung at the point 6. ."
Hero employed a similar argument in his discussion of the ,vheel and
axle. In fact, in reducing the study of these machines to the principle
of circles he was implicitly using the notion of moment. Thus it is
clear, though not explicitly stated, that in his discussion of the axle
Hero understands that a ,veight , can be replaced by an equal force
applied tangentially at
because t4F has the saIne moment as ~2
Fig. 5
2.
PAPPUS'S THEORIES OF THE INCLINED PLANE AND OF THE CENTRE
OF GRAVITY.
Pappus (IVth Century A. D.) appears to he the only geometer of
Antiquity who took up the problem of the motion and equilibrium of
a heavy body on an inclined plane. The proof that we shall analyse
now is taken from Book VIII of his Collections (From among the varied
and delightful problems of mechanics) .
Pappus assumes that a certain effort y is necessary to move a weight
t:J.. on the horizontal plane ltv, and that a power f) is necessary to draw it
1 CARRA DE V AUX'S surmise that Hero is referring to the treatise IIeei 'vywv is
probably correct.
2 Cf. JOUGUET, Lectures de .l\fecanique, ''''01. I, p. 215. Throughout the present
book this treatise will be indicated by the letters L. AI.
:334
THE ORIGINS
IIp the inclined plane /l/~.
He sets out to determine the relation between
y and o.
The weight ex. on the plane It'" has the form of a sphere with centre e.
Pappus reduces the investigation of the equilibrium of this sphere on
K
Fig. 6
th~
inclined plane to the following problem. A balance supported at A
catries the weight oc at e and the weight p which is necessary to keep
it in equilibrium at 1J-theend of the horizontal radius ene The law of
the angular lever, which Pappus borrows from Archimedes' fleel ~vywv
or from Hero's Mechanics, pro"ides the relation
fJ =
ex.
eC
5:.
rj~
On the horizontal plane where the power necessary to move oc is y,
the power necessary to move along fJ will be
Pappus then assumes that the power () that is able to move the
weight ex. on the inclined plane It'" will be the sum of the powers <5 and
y, that is
() == Y
1ALEXANDRIAN SOURCES AND ARABIC ~iANUSCRIPTS
35
.l£vidently the necessity of a power y for pulling the weight ~ on the
~~~~Dtal plane derives from Aristotle's dynamics, in which all unnat-
~.JDotion requires a motive agency. The argllment by which Pappus
introduces the lever fAf] supporting the two ,veights (X and fJ is rather a
natural one, even though it does not lead to a correct evaluation of /3..
The last hypothesis, concerning the addition of band y, the powers
that. are necessary to move fJ and a respectively on the horizontal plane
is, on the other hand, most strange..
However incorrect it may have been, this proof ,"-as destined to
inspire the geometers of the Renaissance.. Guido Ubaldo was to adopt
it and Galileo was to be occupied in demonstrating its falsehood .
Archimedes certainly formulated a precise definition of centre of
gravity, but there is no trace of anything of this kind in those of his
writings that are available to us. Therefore it is of some value to
record the definition which is due to Pappus.
Imagine that a heavy body is suspended
hy an axis rt./3 and let it take up its equili-
brium position. The vertical plane pass-
ing through ct/l ,,' will cut the body into
two parts that are in equilibrium with each
other and which will be hung in such a way,
qn one side of the plane and on the other,
to be equal ~rith respect to their weight. ."
raking another axis a'p' and repeating the
Fig. 7
iiame operation., the second vertical plane
passing through a' P' will certainly cut the
first-if it were parallel to it " each of these two planes would divide
the body into two equal parts which would be at the same time of equal
weight and of unequal weight, which is absurd.. "
Now suspend the body from a point y and draw the vertical y~
through the point of suspension when equilibrium is established. Take
a second point of suspension y' and, in the same way, draw the vertical
,.'3'. The two lines yo, 1"0' necessarily intersect. For if not, through
each of them a plane could be drawn so as to divide the body into two
parts in equilibrium with each other, and in such a way that these two
planes were parallel to each other.. This is impossible.
All lines like yo will therefore intersect at one unique point of the
body that is called the centre of gravity.
Pappus did not distinguish clearly, as Guido Uhaldo was to do in his
Con"mentar_y on ./lrchimedes' two books on the equilibrium of weights (1588)
between '" equiponderant" parts, that is parts that are in equilibrium in
the positions ,\!-hich they occupy, and parts ,vhieh have the same weight.
*836
3.
THE ORIGINS
THE FRAGMENTS ATTRIBUTED TO EUCLID IN ARABIC WRITINGS.
Greek antiquity does not attrihute any work on mechanics to Euclid.
However his name occurs frequently in this connection in the writings
of arabic authors.
Euclid's book on the balance, an arabic manuscript of 970 A. D.
which has been brought to light by Dr. Woepke,l seems to have remain-
ed unkno\vn to the western Middle Ages. This relic of greek science may
be contemporaneous with Euclid and may thus antedate Archimedes.
It contains a geometrical proof of the law of Ie"vers which is independent
of Aristotle's dynamics and which makes explicit appeal to the hypo-
thesis that the effect of a weight P placed at the end of an arm of a lever
is expressed by the product PL. We have had occasion to emphasise
the necessity of this hypothesis in our analysis of Archimedes' proof.
The treatise Liber Euclidis de gravi et levi, often simply called De
ponderoso et levi, ha:5 been known for a long time. It includes a very
precise exposition of aristotelian dynamics which is arranged, after
Euclid's style, in the form of definitions and propositions. The latin
text renders the terms ~{)'VCt.Jl£q; and iaxv;, by which Aristotle meant
" power", as virtus and fortitudo. Bodies that travel equal distances
in the same medium-air or water-in times which are equal to each
other, are said to be equal in t"irtus. Bodies that travel equal distances
in unequal times are of different virtus, and that which takes the shorter
time is said to have the greater virtus. Bodies of the same kind are
those that, volume for volume, are equal in virtus. That which has
the greater virtus is said to be solidius (more dense).
The virtus of hodies of the same kind is proportional to their dimen-
sions ; that is, the bodies fall with velocities which are proportional to
their volume. If two heavy bodies are joined together, tIle velocity
with which the combination will fall will be the sum of the velocities
of the separate bodies.
Duhem has found, in a XIVth Century manuscript,2 four proposi-
tions on questions in statics which complete De ponderoso et levi. This
manuscript contains a theory of the roman balance, and shows that the
fact that the balance is a heavy homogenous cylindel' does not alter the
relation of the weights to each other.
Finally, in a XIIIth Century manuscript, Duhem has unearthed a
text called Liber Euclidis de ponderibus secundunt terminorum circonfe-
rentiam 3 which connects the law of levers ,vith aristotelian dynamics
and also contains a theory of the ronlan balance.
1
2
3
Journal asiatique, Vol. 18, 1851, p. 217.
Bibliotheque Nationale, Paris, latin collection, 1\ls. 10,260.
Ibid., Ms. 16,649.ALEXANDRIAN SOURCES AND ARABIC MANUSCRIPTS
37
BOOK OF CHARISTION.
Charastonis is the latin version of an arabic text due to
ibn Kurrah (836-901). The original greek version
:relna:ms unknown, and the question of whether karaston (in Arabic-
• ·I«I,rsJ~ftn) refers merely to the roman balance or to the name of the greek
f!e.)m,etf~r Charistion (a contemporary of Philon of Byzantium in the
B. C.) has been the subject of much scholarly debate.
shall
Duhem 1 in summarising the theory of the roman
balance which is found in Liber Charastonis.
2e(J~me~ter Thitbit
b
d
u
9
a
e
Fig. 8
A heavy homogeneous cylindrical beam a,b whose arms ag and bg
~:re.
unequal may he maintained in a horizontal position by means of a
If bd is the amount
~y which the longer arm exceeds the shorter arm and u is the centre of
/6·a, the weight e ,viII be to bd as gu is to ga. If p is the total weight of
It the beam
iw'eight e hung from the end of the shorter arm ag.
db
e -- p - -,....
---
2ga
If this weight were known it could be represented exactly by a
hung from the shorter arm, and the karaston arranged in this
wfl.ycould be treated as a weightless heam.
~. We must also mention, as one of the sources of statics, the treatise
Canonio,2 a latin translation of a greek text ,vhich adds nothing
essential to Li ber Charastonis.
~eAl'C .. pan
1
2
O. S., Vol. I, p. 90.
Bibliotheque Nationale, Paris, latin collection, Ms. 7378 A.CHAPTER THREE
THE Xlllth CENTURY
THE SCHOOL OF JORDANUS
1.
]ORDANUS OF NEMORE AND" GRAVITAS SECUNDUM SITUM. "
The Middle Ages had access to the Problems of Mechanics and to the
works of Aristotle. They had also inherited the fragments attributed
to Euclid-with the exception of the Book on the Balance-as well as
the Liber ClJ,arastonis from arabic learning. They had no knowledge
of Archimedes, Hero of Alexandria and Pappus.
In spite of the researches of the scholars, the personality of J ordanu8
remains mysterious. At least three XllIth Century manuscripts on
statics; have been attributed to him, although these are clearly in the
style qf different authors. Neither Jordanus's nationality nor the pe-
riod id which he lived is known with any certainty. Daunou believes
him t9 have lived in Germany about 1050, Chasles associates him with
the ~IIIth Century while Curtze places him about 1220 under the name
of Jbrdanus Saxo. Michaud has identified him with Raimond Jordan,
provost of the church of Uzes in 1381 which is clearly too late. With
Montucla, we shall here adopt the intermediate vie,v that associates
Jordanus of Nemore with the XIIlth Century.
Like Duhem, we shall follow the Elementa J ordani super demonstra-
tionem ponderis. 1 This work comprised seven axioms or definitions
foll().wed by nine propositions. The essential originality of Jordanus lay
in the systematic use, in his study of the motion of heavy bodies, of the
effective path in a vertical direction as a measure of the effect of a weight,
which wa-s usually placed at the end of a lever and described a circle in
consequence. Thus his statics stems, implicitly, from the principle of
virtual work. The word work, taken in the modern sense., is to be con-
1 Bibliotheque Nationale, Paris, Ms. 10,252, dated 14.64.
There also exists an in·
complete manuscript of the same work, dating from the XIlIth Century, in the
Biblio·
theque Mazarine, Ms. 3642.39
THE XI11th CENTITRY
trasted with the word velocity and with the concept of virtual velocities
l\rhich may he traced in the arguments of Problems of Mechanics. Of
course lordanus never used the word" work" itself.. He considered
the heaviness of a particle relative to its situation (gravitas secundum
situm) without making clear the relation that exists between this quan-
tity and the heaviness in the strict sense.
Jordanus formulated his principle in a picturesque Latin which merits
quotation.
" Omnis ponderosi motam esse ad lnedium, virtutemque ipsius poten-
tiam ad inferiora tendendi et motui contrario resistendi.
" Gravius esse in descendendo quando ejusdem motus ad medium rectior.
" Secundum situm gravius, quando in eodem situ minus obliquus est
descensus.
" Obliquiorem autem descensum in eadem quantitate minus capere de
directo. "
Or-
" The motion of all heavy things is towards the centre,! its strength
heing the power ,vhich it has of tending do,vnwards and of resisting a
contrary motion.
,,4 A moving body is the heavier in its descent as its motion towards
the centre is the more direct.
'" A body is the heavier because of its situation as, in that situation,
descent is the less oblique.
,~ A more oblique descent is one that, for the same path, takes less
of the direct. "
Thus a certain weight placed at b,
at the end of the lever cb, has a smaller
graVity secundum situm than the same
~~~ght has when it is at a, at the end
of the horizontal radius ca. Indeed, on
a....- - - -.. c
the circumference of the circle with
centre c and radius ca == cb, if the
- - --~z'
:b,ody falls from b to It along the arc
I
bii the effective path in a vertical
I
direction is b' h'. On the other hand
if. ,the body starts from a and falls
Fig. 9
along an ~ arc liZ., ","hich is equal to
the arc bh, the effective vertical path
is cz' and is greater than b' h'. Thus the descent bh, equal to the
descent liZ., is more oblique than that and takes less of the direct.
,
1
ITnderstood as the common centre of all heavy things in . ' . \'ristotle's sense.40
THE ORIGINS
This idea led J ordanus to a proof of the rule of the equilibrium of the
straight lever whose originality cannot be contested.
d
b
" Let acb he the hearn, a and b the weights that it carries, and suppose
that the relation of b to a is the same as that of ca to cb. I maintain
that this rule will not change its place. Indeed, if the arm supporting b
falls and the heam takes up the position dee, the weight b will descend
by he ~nd a will rise by fd. If a weight equal to the weight b is placed
at 1, at a distance such that cl == eb, this will rise in the motion by
gm === he. But it is clear that dfis to mg as the weight I is to the weight a.
Consequently, what is sufficient to bring a to d will be sufficient to bring
I to m. But we have shown that band 1 counterbalance each other exact-
~y, so that the supposed motion is impossible. This will also he true
jof the inverse motion. "
Duhem writes in this connection 1 _
'" Underlying this demonstration of Jordanus the following principle
is clearly evident-that which can lift a weight to a certain height can
also lift a weight which is k times as
8
c
great to a height which is k times less.
This principle is then the same as that
which Descartes took as a basis for his
complete theory of statics and which,
thanks to John Bernoulli, became the
principle of virtual work. "
Jordanus was less fortunate "\\Then he
turned his attention to the angular lever.
He considered a lever ac! carrying equal
f
weights at a and f which were placed in
Fig. 11
such positions that ac == ef. Jordanus
was of the opinion that, under these
1
o.
S., Vol. I, p. 121.THE Xlllth CENTURY
·41
would dominate f. He arrived at this conclusion by con-
equal arcs ;;1 and ;f. It is clear that the "Co direct " taken
a is greater than the " direct " taken by the weight f.
conclusion is obtained hecause, since the linkages are
two displacements;;] and j;;" arc not simultaneously possible.
thus misunderstood the idea of moment.
as the XIIIth Century the Elementa Jordani were generally
copyi:;ts with lJe Canonio to form the Liber Euclidis de
This artificial associations and this imaginative titles
despair of the scholars and it has needed all the learning of
................,.,. . . . . to elucidate them.
Every truly novel idea evokes a reaction. The Elementa Jordani
did not provide an exception to this rule. In the XIIIth Century a
eritie wrote a commentary of J ordanus which Duhem calls the Peripa-
tetic Con~n~entary.2 This author at every turn invokes the authority of
Aristotle and has scruples ahout applying the gravitas secundum situm
to .8 motionless point-in modern language, about making appeal to
amtual displacement. It does not appease his conscience to consider
that rest is the end of motion. "" The scientific value of the Commen-
tary is nothing," declares Duhem. 3 "But its influence did not
disappear for a very long time, and even the great geometers Tarta-
gJia, Guido Uhaldo and Mersenne had not entirely freed themselves
ftom it. "
2}~ THE ANONYMOUS AUTHOR OF "LIBER JORDANI DE RATIONE PON..
,
DERIS .. "
TIlE ANGULAR LEVER.
THE INCLINED Pl,ANE.
We now come to an especially noteworthy work which figures in the
same manuscript as the Peripatetic Commentary under the title Liber
Jii,i!a,ni de ratione ponderis, and which did not remain unknown in the
Renaissance. l'artaglia sent it to Curtius Trojanus who puhlished it
in 1565. This work supercedes and rectifies the Elementa Jordani on
m~ny important points. ...J\ll the same, it is hased on the same principle
of gravitas secundum situm.
Duhem, ,\Tho brought this manuscript to light, terms the anonymous
author a H Precursor of Leonardo da Vinci." Indeed, in many respects
this "precursor surpassed Leonardo, who, for example, spent himself
in fruitless efforts to evaluate the apparent weight of a hody on an
inclined plane.. It seems more natural to simply speak of an anonymous
1
2
3
Bibliotheque Nationale, Paris, latin collection., Mss. 7310 and 10.,260.
Ibid., 1\'15. 7378 A.
O. S.') 'Tol. I, p. ]34.42
THE ORIGINS
author of the XIIIth Century, a disciple of Jordanus who had out..
stripped his master.
In connection with the hent lever this author corrected Jordanus's
error. As before, let a lever acf carry equal ,veights at a and f and
be placed in such a position that aa' == if'.
c
I
It is impossible that the weight a should dominate the weight f.
!For if two arcs ;;}", fl, are considered on the two circles drawn through
a and f and corresponding to equal angles -;;J" and fii; the descent
of a along rh necessitates that the equal weight at f should rise through
a distance In which is greater than rh. This is impossible.
In the same \vay it can be seen that f will not dominate a. For
if the arcs j; and ~ correspond to equal
c
anglesft; and;;;;;;' the descent ofjalong
tx makes it necessary that the equal
I
weight placed at a should rise by pm,
a
~------~ J
a
\vhich is greater than tx. This is impos-
I
sible. Therefore there is equilibrium in
I
the position considered, in which aa' if'.
J
The anonymous author generalised
b l - b
this result to an angular balance
I
Fig. 13
carrying unequal weights at a and b,43
THE XIIIth CENTURY
and obtained the result that in equilibrium it is necessary that the
distances aa' and bb' from a and b to the vertical drawn through the
point of support, c, are in inverse ratio to the weights a and b..
We see that this author knew and used the notion of moment..
Elsewhere he wrote on this subject., '" If a load is lifted and the
length of its 81lpport is known, it can be determined how much
this load weighs in all positions. The weight of the load carried at
e by the support be will he to the weight carried at f by fb as el is
to fr or as pb is to xb. .A. ,veight placed at e, at the end of the
lever be, will weigh as if it were at
u on the lever bj: "
Thus the idea of gravitas secundum
situm, which Jordanu8 had used qua-
litatively, became precise.
Our anonymous author also con-
cerned himself with the stability of
the halance, and rectified certain
errors which were contained in the
relevant parts of Problems of Mecha-
nics.
C .....
p - -.. b
More than this, he resolved the
Fig. 14
problem of the equilibrium of a heavy
h~dy on an inclined plane, a problem
wbich had eluded the wisdom of the greek and alexandrian geometers..
, In order that this may be done, it is first observed that the gravitas
stcull,dum situm of a weight on an inclined plane is independent of
its position on the plane. The author then attempts a comparison of
, the value that that gravity takes on differently inclined planes. We
shall quote from Duhem's translation of this same Xlllth Century
manuscript.
'" If two ,veights descend by differently inclined paths, and if they
are directly proportional to the declinations, they ,,,ill be of the sanle
strength in their descent..
,<. Let ab be a horizontal and bd, a vertical.
Suppose that two
oblique lines da and de fallon one side and on the other of bd, and
that de has the greater relative ohliquity.. By the relation of the ohli-
quities I understand the relation of the declinations and not the relation
of the angles; this means the relation of the lengths of the named
lines counted as far as their intersection with the horizontal, in such
a way that they take simiiarily of the direct.
" In the second place, let e and h be the weights placed on dc and
da respectively, aDd suppose that the ",-eight e is to the ,veight h as
,
-...r-----.. .44
THE ORIGINS
d
-----,.,ja...----......---------..
k ....
C
Fig. 15
de is to da. I maintain that in such a situation the two weights will
have th.e same strength.
" Indeed, let dk be a line having the same obliquity as de and,
on that line, let there be a weight g which is equal to e.
" Suppose that the weight e should descend to I, if that is possible.,
and that it should draw the weight h to m. (It is clear that the author
imagines the two weights to he connected by a thread which passes
over a pulley at d.) Take gn equal to hm, and consequently equal
to el. Draw a perpendicular to db which passes through g and h,
say ghy. Drop a perpendicular It from the point I onto db. Then
drop [the perpendiculars] nr, mx, and ez. The relation of nr to ng is
tha~ of dy to dg and also that of db to dk. Therefore mx is to nr as
dk i$ to da ; that is to say, as the weight g is to the weight h. But as e
is nJ)t able to pull g up to n (nr ::=: ez), it is no better ahle to pull h up
to /m. The weights therefore remain in equilibrium. "
This demonstration, which leads to the correct law of the apparent
heaviness of a body on an inclined plane, was directly inspired hy that
"of Jordanus concerning the equilibrium of a straight lever. Like
that, it, implicitly proceeds according to the principle of virtual work.
We shall now give some indication of the ideas on dynamics which
were used by the author of Liber Jordani de ratione ponderis.
The environment's resistance to the motion of a hody depends
on the shape of the body, which penetrates the environment the hetter
as its shape is the more acute and its figure the more smooth. It
depen.ds~ in the second place, on the density of the fluid traversed. All
media are compressihle; the lower strata, compressed by the upper
ones., are the denser and those which hinder motions more. At the
front of the moving body will be a part of the medium cQmpressed
on, and sticking to it. But the other parts of the medium, which
are displaced by the mo
ligand effect
Many hormones cannot pass through the
plasma membrane; instead, they interact with
C
CONTABI LITATEA – FORMĂ
Ă DE BAZĂ
Ă A EVIDE
ENȚEI EC ONOMICE
E
1.1
1. Obiective e
La a sfârşitul ac cestei unităţ ţi de învăţar re studenţii vor fi capab bili să:
 define ească noțiun
nea de conta abilitate;
 înțelea agă necesi itatea organ nizării con ntabilității la nivelul fiecărei
unităț ți patrimonia ale;

înțelea agă rolul pe care îl are conta abilitatea în n cadrul activității
a
desfăș șurate de un
nitățile patri imoniale;

înțelea agă modul de
d organizar re a contabil lității la nive el microeco onomic.
1.2. Definir rea, necesit tatea și rolu
ul contabili ității
C
Contabilitat
tea a exista at din cele e mai vech hi timpuri. C.G. Dum
mitrescu, în „Istoria
contabil lității”, arăta a că grecii au
a împrumu utat tehnica evidenței co ontabile de la egipteni, iar de la
ei au pr reluat-o rom
manii. Dar se pare ca evidențele contabile sunt
s
mult mai
m vechi în n istoria
omeniri ii. Contabil litatea în pa artidă dubl ă s-a născu ut ca urma are a practi
cii contabil lilor din
Veneția a și Genova a. În anul 1 494, Luca Paciolo des scrie contab
bilitatea în partidă
p
dub blă într-o
lucrare de matema atică și geom
metrie. Dup
pă apariția acestei luc crări, aplicar rea contabi ilității în
partidă dublă s-a ră ăspândit și în
î alte țări a le Europei.
A
Astăzi,
con
ntabilităţii îi i revin sarci ini din ce în n ce mai gr rele. Ea cau
ută să-şi dep păşească
limitele , fiind pusă ă în situaţia de a descrie e organizaţi ii din ce în ce
c mai comp
plexe care operează
o
într-un m
mediu econ nomic şi soc cial în contin
nuă mişcare e şi transfor rmare.
Princip
palele aspec cte  ca ins strument de descriere, de
d modelare e a întreprin nderilor;
sub car re trebuie  ca ins strument de prelucrare a informaţii ilor;
studiată ă contabilit tatea  ca pr ractică sau u „joc soci ial”, înscri să într-o
r reţea de
(după N.
N Feleagă) )
restric cţii regleme entare mai mult
m sau ma i puţin stric cte.
10Contabilitatea poate fi considerată drept o artă, o tehnică sau o ştiinţă, dar
indiferent de
cum am privi-o, contabilitatea este „un joc social” ce are drept finalitate
reprezentarea unei
realităţi care este entitatea.
Contabilitatea studiază acele laturi ale reproducţiei sociale care se pot exprima
în
etalon bănesc. Ea urmăreşte existenţa şi dinamica patrimoniului agenţilor
economici,
procesele economice, pe care aceştia le organizează, stabilind şi înregistrând
rezultatele
financiare finale.
1.3. Organizarea evidenţei contabile la nivel microeconomic
Conform Legii Contabilităţii nr. 82/1991, întreprinderile au obligaţia să
organizeze şi
să conducă contabilitate proprie, în limba română şi în moneda naţională.
Organizarea
contabilităţii reprezintă deci, nu numai o necesitate, aşa cum am arătat anterior,
dar şi o
obligaţie impusă prin reglementările legale în vigoare.
De altfel, entitatea ca sistem complex economico-social şi administrativ-
organizatoric,
îndeplineşte o serie de funcţii, în cadrul cărora un rol esenţial îl are funcţia
financiar-
contabilă.
Pentru îndeplinirea acestei funcţii şi în acelaşi timp pentru respectarea legii, în
cadrul
entității se organizează şi funcţionează un compartiment specializat, financiar-
contabil. În
cadrul acestui compartiment lucrează persoane cu studii de specialitate (medii şi
superioare),
având atribuţii distincte în domeniul evidenţei contabile operative şi generale.
Compartimentul financiar-contabil se subordonează contabilului şef, care are studii
superioare
în finanţe-contabilitate.
Aici se consemnează zilnic şi lunar, toate operaţiunile economico-financiare ce au
loc
în entitate, respectiv: cumpărări, vânzări, consumuri, salarii, încasări, plăţi
etc., conducând în
final la determinarea rezultatului activităţii şi la întocmirea situaţiilor
financiare anuale de
sinteză şi raportare contabilă.
Luna calendaristică poartă denumirea de perioadă de gestiune, iar anul
calendaristic,
de exerciţiu financiar.
În baza datelor furnizate de evidenţa contabilă, se pot efectua analize economico-
financiare privind corelarea resurselor alocate cu rezultatele obţinute, se pot
calcula diverşi
indicatori şi se poate determina evoluţia diverselor fenomene în timp, cu factorii
pozitivi şi
negativi care le-au generat.
11În cadrul entității, contabilitatea se organizează pe două circuite paralele:
contabilitate
financiară şi contabilitate de gestiune.
 este reglementată prin norme unitare;
 oferă o viziune globală asupra activităţii;
 are un obiectiv financiar – reflectarea imaginii fidele a
patrimoniului;
Contabilitatea financiară  generează fluxuri de informaţii şi documente externe;
 aplică reguli normative;
 oferă date utilizatorilor externi (furnizori, clienţi, bănci,
investitori, organe de control etc.);
 se referă la perioade încheiate (lună, an).
 se lasă la latitudinea fiecărei entități;
 oferă o viziune detaliată asupra activităţii;
 are un obiectiv economic constând în supravegherea şi
controlul activităţii prin intermediul costurilor;
Contabilitatea  generează fluxuri de informaţii interne;
de gestiune  aplică reguli stabilite în cadrul entității;
 oferă date conducerii entității;
 se referă la prezent şi viitor (previziuni);
 clasifică cheltuielile după locul de realizare şi destinaţia
lor.
12După parcurgerea acestei unităţi de învăţare trebuie să reţineţi:

Ce este contabilitatea şi care este rolul acesteia în cadrul unei
entităţi.

Care sunt circuitele de organizare ale contabilităţii în cadrul
unei entităţi.
 Ce este contabilitatea financiară şi prin ce se caracterizează.
13UNITATE
EA DE ÎNV
VĂȚARE 2
OBIEC
CTUL ȘI METODA
M
C
CONTABIL
LITĂȚII
2.1
1. Obiective e
 identi fice obiectu ul de studiu al contabili ității;
 identi fice proced deele ce apar rțin metode i contabilită ății;
 identi fice și să în nțeleagă rolu ul principiil or contabili ității;
 identi fice funcții ile contabil lității și ne ecesitatea ap
plicării ace estora în
contab bilitatea fin anciară și în n contabilita atea de gest tiune.
2 Obiect tul contabil lității
2.2.
C orice di sciplină știi ințifică, con ntabilitatea reprezintă simultan
Ca
s
o teorie
t
și me etodă. În
calitatea a sa de teor rie științific că, contabil itatea repre ezintă un sis stem
de pri incipii și cu unoștințe
care ex xplică și inf formează, iar
i ca meto
odă sau teh hnică, un an nsamblu co oerent de procedee,
p
instrume ente prin ca are se observ vă și înregis strează resu ursele econo omice
ale so ocietății, sep parate ca
utilități patrimonial le.
Concep
pții cu
1. Concep pția admin nistrativă consideră că obiectu ul contabil lității îl
privire la constit tuie reflect area și con ntrolul, în expresie valorică,
v
a faptelor
delimita area admini istrative, în n vederea sp prijinirii ma anagementu ului,
pentru a obține
și defin
nirea cu un minim de eforturi (ch heltuieli) un n maxim de e efecte eco onomice
obiectu
ului (rezult tate-venituri i).
contabi ilității
2. Concep pția juridic că consider ră că obiec ctul contab bilității îl formează
f
patrim
moniul unui i subiect de
d drept (a agent econo
omic), prin n prisma
relațiil or juridice e, adică a drepturil lor și obl ligațiilor p pecuniare
(mater riale) ale unei
u
persoa ane fizice sau juridic ce, în core elație cu
obiecte ele, adică cu
u bunurile și
ș valorile co orespunzăto oare.
3. Concep pția
econ omică
sau u
financia
ară
consid
deră
că
obiectul
contab bilității îl co
onstituie cir rcuitul capit talului, priv vit atât sub aspectul
destina ației lui, câ ât și sub for rma modulu ui de dobân ndire, respe ectiv sub
forma de capital propriu
p
și ca apital străin. .
16 Obiectul contabilității este reprezentat de reflectarea în expresie bănească
a bunurilor mobile și imobile, inclusiv solul, bogățiile naturale,
zăcămintele și alte bunuri cu potențial economic, disponibilitățile bănești,
Concluzie
titlurile de valoare, drepturile și obligațiile agenților economici, precum
și mișcările și modificările intervenite în urma operațiunilor patrimoniale
efectuate, cheltuielile, veniturile și rezultatele obținute de aceștia.
 Obiectivul fundamental al contabilității îl reprezintă furnizarea de
informații utile luării deciziilor menite să asigure o imagine fidelă a
patrimoniului, situației financiare și a rezultatelor obținute.
Din această definiție rezultă că patrimoniul presupune existența a două condiții de
bază: un titular de patrimoniu (subiect de drept) pe de o parte, iar pe de altă
parte bunurile și
valorile economice, privite ca obiecte de drepturi și obligații, precum și
titularul de patrimoniu.
Structura de ansamblu a patrimoniului este următoarea:
PATRIMONIU
BUNURI ECONOMICE
PATRIMONIU PROPRIU
(DREPTURI)
PATRIMONIU STRĂIN
(OBLIGAȚII)
Bunurile economice se concretizează în bunuri materiale (terenuri, clădiri,
utilaje,
mijloace de transport, stocuri de materii prime etc.) şi nemateriale (proiecte,
programe
informatice etc.) formând averea entității.
Relaţiile de drepturi se referă la faptul că entitatea îşi procură o parte din
avere din
surse proprii, iar bunurile procurate îi aparţin de drept.
Relaţiile de obligaţii se referă la faptul că entitatea îşi procură o parte din
avere din
surse împrumutate, fiind obligată să restituie terţilor contravaloarea bunurilor
procurate.
Contabilitatea se ocupă cu reflectarea în expresie valorică a patrimoniului, ea
înregistrează circuitul elementelor patrimoniale în condiţii concrete de timp şi
spaţiu,
calculează mărimea acestor elemente şi reflectă mişcarea patrimoniului prin
operaţiuni de
intrări şi ieşiri 1 .
1
Ghe. Talaghir, Ghe. Negoescu – Contabilitatea pe înţelesul tuturor. Editura All,
Bucureşti, 1998.
17Contabilitatea studiază modul de gestionare a patrimoniului, fundamentează
deciziile referitoare la finanţarea şi utilizarea elementelor patrimoniale,
controlează realizarea
deciziilor şi stabileşte răspunderi privind integritatea şi dezvoltarea
patrimoniului.
De asemenea, contabilitatea studiază echilibrul global al patrimoniului, prin
respectarea ecuaţiei patrimoniale:
BUNURI ECONOMICE = DREPTURI + OBLIGAȚII
cu derivatele sale:
Drepturi = Bunuri economice – Obligaţii
şi
Obligaţii = Bunuri economice – Drepturi
În consecinţă, entitatea reprezintă o unitate patrimonială, al cărei patrimoniu
poate fi
privit sub dublu aspect: al mijloacelor economice (bunurile/averea) şi al surselor
de
procurare a acestor mijloace (capitalul) proprii şi străine.
Mijloacele economice definesc activul patrimonial, iar sursele definesc pasivul
patrimonial.
În derularea operaţiunilor economico-financiare, apar şi o serie de procese
economice, sub forma veniturilor şi cheltuielilor, care ajută la înregistrarea
creşterii sau
diminuării patrimoniului.
În acest context, ecuaţia patrimonială de mai sus, devine:
AVERE = CAPITAL
182.3. Metoda contabilității
Datorită complexității obiectului de studiu, metoda contabilității reunește mai
multe
procedee tehnice de lucru.
Un ansamblu de procedee aflate într-o strânsă corelație și
Metoda intercondiționare ca un tot unitar, în vederea stabilirii normelor și
contabilității principiilor cu caracter special pe care se fundamentează
contabilitatea
și cu ajutorul cărora cercetează starea și mișcarea elementelor
patrimoniale ale unităților patrimoniale.
O trasătură caracteristică a metodei contabilității este aceea a folosirii unor
procedee
care să permită înregistrarea numerică, cifrică, a existenței și mișcării
patrimoniului unităților
economice și sociale în expresie valorică. Generalizând, se poate spune că metoda
contabilității reprezintă totalitatea procedeelor interdependente, pe care le
folosește aceasta în
scopul cunoașterii situației patrimoniului și a rezultatelor obținute.
1. procedee comune tuturor științelor;
Procedee utilizate 2. procedee specifice metodei contabilității;
de contabilitate 3. procedee ale metodei contabilității, comune și altor discipline
economice.
Procedee comune Observația – este faza inițială a cercetării obiectului de studiu
al
tuturor științelor oricărei științe și este utilizată pentru cunoașterea
operațiilor economice
care se pot exprima valoric și pe care le reflectă cifric, numeric, cu
ajutorul procedeelor sale specifice.
Raționamentul – se aplică de metoda contabilității, pentru că pe bază
de judecăți logice, pornind de la fenomenele și procesele economice
care intră în obiectul său de studiu, să ajungă la concluzii noi (ex.:
activul este egal cu pasivul, pentru că între mijloacele economice și
sursele de finanțare a acestora există o egalitate perfectă).
Comparația – se folosește de metoda contabilității prin alăturarea a
două sau mai multe fenomene și procese economice care se pot exprima
19valoric, cu scopul de a stabili asemănările și deosebirile dintre ele, ca
astfel să se tragă o serie de concluzii. Se folosește frecvent pentru a se
compara veniturile și cheltuielile pe baza cărora se stabilesc rezultatele
finale.
Clasificarea – acțiunea de împărțire, distribuire, repartizare sistematică
pe clase sau într-o anumită ordine a obiectelor în funcție de asemănările
și deosebirile dintre ele. Asemănările le apropie și le încadrează în
aceeași clasă, iar deosebirile le diferențiază și le distribuie în clase
diferite.
Analiza – procedeu științific de cercetare a unui întreg, a unui fenomen
care se bazează pe examinarea fiecărui element component în parte.
Sinteza – ca procedeu științific de cercetare a fenomenelor se bazează
pe trecerea de la particular la general, de la simplu la compus pentru a
se ajunge la generalizare.
Procedee specifice Bilanțul – stă la baza dublei reprezentări a patrimoniului în
metodei contabilitate. El furnizează informații generale privitoare la situația
contabilității economică și financiară a unei entități, dar reflectă și la
relațiile ei
economice cu alte entități, fiind completat de o serie de situații anexă
prin care se explică și se detaliază anumite laturi ale activității
economico-financiare ale societății.
Contul – se deschide în contabilitatea curentă pentru reflectarea
fiecărui element din patrimoniu. Contabilitatea dispune de un sistem
de conturi în care reflectarea operațiilor economice rezultate din
mișcarea elementelor patrimoniale are la bază dubla înregistrare.
Balanța de verificare – asigură în contabilitatea dublei reprezentări și
a dublei înregistrări, garanția exactității înregistrărilor efectuate în
conturi. Datele balanței de verificare stau la baza întocmirii bilanțului.
Balanța de verificare îndeplinește atât o funcție de control, cât și o
funcție economică, constituind puntea de legătură între cont și bilanț.
20Procedeele metodei Documentația – orice operație economică și financiară
referitoare la
contabilității comune existența și mișcarea elementelor patrimoniale trebuie să fie
și altor discipline consemnată în documente care fac dovada înfăptuirii lor.
economice Evaluarea – procedeul prin care datele contabilității sunt reprezentate
printr-o singură unitate de măsură, creând posibilitatea centralizării
lor cu ajutorul balanțelor de verificare și generalizarea cu ajutorul
bilanțului. Evaluarea constă în transformarea unităților naturale în
unități monetare cu ajutorul prețurilor.
Calculația – este strâns legată de evaluare ca procedeu al metodei
contabilității. Acest procedeu își găsește aplicarea cea mai largă în
domeniul calculației costurilor de producție.
Inventarierea – se folosește pentru a se cunoaște situația reală a
patrimoniului reflectat în contabilitate, și trebuie să se verifice
existența faptică a tuturor elementelor sale, în scopul descoperirii
neconcordanțelor dintre datele înregistrate în conturi și realitatea de
pe teren.
2.4. Principiile contabilității
Pentru a se oferi o imagine fidelă a patrimoniului, a situației financiare și a
rezultatelor
obținute de către entitate, trebuie respectate cu bună credință regulile privind
evaluarea
patrimoniului și celelalte norme și principii contabile.
Principiul  nu sunt admise supraevaluarea elementelor de activ și a
prudenței veniturilor, respectiv subevaluarea elementelor de pasiv și a
cheltuielilor, ținând cont de deprecierile, riscurile și pierderile
posibile generate de desfășurarea activității unității;
 prudența presupune anticiparea efectelor unor acțiuni și în special
a transferului de proprietate cu efecte posibile asupra exercițiului
sau a celor parcurse deja, întrucât asupra lor nu se mai poate
interveni din punct de vedere contabil;
 la încheierea fiecărui exercițiu financiar se contabilizează
datoriile și pierderile probabile.
21Principiul
 continuitatea aplicării regulilor și normelor privind evaluarea,
permanenței înregistrarea
în
contabilitate
și
prezentarea
elementelor
metodelor patrimoniale și a rezultatelor asigurând comparabilitatea în timp a
informațiilor contabile;
 asigură aplicarea pentru aceleași elemente, stucturi, domenii de
activitate, a acelorași metode de la un exercițiu la altul, excluzând
schimbarea metodelor în cursul exercițiului;
 modificarea metodelor de la un an la altul trebuie să fie
determinată de o profundă motivație (modificarea unor acte
normative, stabilirea unor reguli generale de evaluare noi etc.).
Principiul
 presupune că unitatea patrimonială își continuă în mod normal
continuității funcționarea într-un viitor previzibil, fără a fi în stare de
lichidare
activității sau de reducere sensibilă a activității;
 atunci când funcționarea este delimitată în timp sunt menționate
datele de începere și de încetare a activității;
 permite fixarea responsabilității entității în raport cu terțe
persoane, inclusiv cu bugetul statului.
Principiul
 presupune delimitarea în timp a cheltuielilor și veniturilor
independenței aferente activității unității patrimoniale pe măsura angajării
exercițiului acestora și trecerii acestora la rezultatul exercițiului la care se
referă;
 independența exercițiului trebuie înțeleasă prin aceea că nu se mai
poate interveni asupra bilanțului după perioada de raportare
(aprobarea consiliului de administrație, înregistrarea la organele
financiare etc.).
Principiul
intangibilității
 bilanțul de deschiderea unui exercițiu trebuie să corespundă cu
bilanțul de închidere al exercițiului precedent.
bilanțului de
deschidere
22Principiul
necompensării
 elementele de activ și de pasiv trebuie să fie evaluate și
înregistrate în contabilitate separat, nefiind admise compensarea
între posturile de activ și de pasiv ale bilațului;
 necompensarea trebuie înțeleasă în sensul că pentru fiecare
element patrimonial cu substață materială, pentru orice resursă
care reflectă drepturi și obligații trebuie să fie deschis, în
contabilitate câte un cont nou;
 necompensarea nu trebuie confundată cu rectificarea prevăzută în
scopul reflectării valorii rămase;
 valorile rectificative sunt evidențiate distinct chiar dacă în cazul
unor elemente patrimoniale înregistrarea în bilanț se face la
valoarea considerată netă sau rămasă (imobilizări rectificate pe
calea amortizării, materiale de natura obiectelor de inventar
rectificate pe calea uzurii sau deprecierilor rectificate pe seama
provizioanelor etc.).
2.5. Funcțiile contabilității
Contabilitatea
financiară
 Funcția de înregistrare completă a tranzacțiilor entității în scopul
determinării periodice a situației patrimoniale și a rezultatului global;
 Funcția de comunicare financiară externă (informarea terților);
 Funcția de instrument de verificare și de probă judiciară și fiscală;
 Funcția de instrument de gestiune a entității;
 Funcția de furnizare a informațiilor necesare realizării sintezelor
macroeconomice;
 Funcția de informare pentru analize financiare.
Contabilitatea
internă de gestiune
 Funcția de determinare a costurilor pe produse, lucrări și sectoare
de activitate;
 Funcția de determinare a diferitelor marje și a rezultatelor
analitice pe produse și activități;
 Funcția de generalizare și furnizare a informațiilor destinate
elaborării bugetelor și costurilor previzionate;
23 Funcția de generalizare și furnizare a informațiilor destinate
actualizării indicatorilor de gestiune din structura tabloului de
bord al entității;
 Funcția de generalizare și furnizare a informațiilor destinate
măsurării performanțelor la nivelul sectoarelor și pe produse,
lucrări și servicii.
După parcurgerea acestei unităţi de învăţare trebuie să reţineţi:
 Care este obiectul de studiu al contabilității.
 Care sunt procedeele specifice metodei contabilității.
 Care sunt principiile contabilității.
 Care sunt funcțiile contabilității și necesitatea aplicării acestora
în contabilitatea financiară și în contabilitatea de gestiune.
24UNITATEA DE ÎNVĂȚARE 3
ACTIVUL PATRIMONIAL
3.1. Obiective
La sfârşitul acestei unităţi de învăţare studenţii vor fi capabili să:
 definească noțiunea de active patrimoniale;
 identifice și să descrie elementele componente ale activelor
imobilizate;

identifice și să descrie elementele componente ale activelor
circulante.
3.2. Activul patrimonial
Este format din totalitatea mijloacelor economice destinate bunei desfăşurări a
activităţii. Elementele patrimoniale de activ se pot clasifica: după modul de
valorificare şi după
gradul de lichiditate. Se observă că, după ambele criterii se conturează două mari
categorii de
active patrimoniale: active imobilizate şi active circulante.
I. Activele imobilizate
- active fixe ce se consumă şi se valorifică în mod treptat, pe
parcursul mai multor cicluri de exploatare, având un grad de
lichiditate scăzut.
În funcție de natura lor, ele cuprind următoarele grupe:
1) Active imobilizate
necorporale
- sunt active fixe nemateriale, reprezentate prin documente juridice
sau comerciale ce conferă entității anumite drepturi.
- sunt active fixe materiale.
corporale
financiare
- sunt active fixe de natura investiţiilor financiare în cadrul altor
întreprinderi, efectuate pe termen mediu şi lung, în scopul
obţinerii de venituri.
1) Activele imobilizate necorporale cuprind:a) Cheltuielile de
constituire
- sunt cheltuielile efectuate de entitate la înfiinţarea ei: cheltuieli
de înscriere la Registrul Comerţului, taxe de publicare la
Monitorul Oficial, cheltuieli cu emisiunea de acţiuni şi părţi
sociale etc.
b) Cheltuielile de
cercetare-dezvoltare
- sunt cheltuieli efectuate pentru achiziţionarea sau realizarea în
producţie proprie a unor proiecte de cercetare ce urmează a fi
aplicate în procesul de producţie, în scopul obţinerii de profit.
c) Concesiunile
- sunt convenţii prin care entitatea dobândeşte, în schimbul unei
sume, dreptul de a exploata pe o anumită perioadă, bunuri
proprietate a statului (terenuri, clădiri etc.).
d) Brevetele
- sunt documentele prin care i se recunoaşte unei persoane dreptul
de a exploata exclusiv un produs al cărui autor este.
e) Licenţele de
fabricaţie
f) Mărcile de fabrică
- sunt drepturi câştigate de entitate, de a exploata un brevet, prin
cumpărarea acestuia.
- sunt sume investite de entitate pentru ca produsele sale să se
deosebească de produse similare existente pe piaţă.
g) Fondul comercial
- reprezintă dreptul suplimentar cuvenit entității, peste valoarea
bunurilor materiale, datorită existenţei unor condiţii deosebite
precum: vadul comercial, clientela etc.
h) Programele
- reprezintă partea soft a echipamentului electronic, ce permite
informatice efectuarea diverselor operaţiuni cu ajutorul tehnicii de calcul.
i) Alte imobilizări - cuprind cheltuieli ocazionate de achiziţionarea sau
elaborarea
necorporale
j) Imobilizările
necorporale în curs
programelor informatice şi alte imobilizări necorporale.
- reprezintă costul de producţie sau de achiziţie a imobilizărilor
necorporale neterminate până la sfârşitul exerciţiului financiar.
Datorită utilizării lor pe durată îndelungată, imobilizările necorporale se supun
uzurii în
timp. Expresia valorică a uzurii imobilizărilor se înregistrează în contabilitate
sub forma
amortizării.
2) Activele imobilizate corporale cuprind:
a) Terenurile
- în funcție de destinație se clasifică în: terenuri agri
cell surface receptors. Their effects are direct
and very rapid. With ligand-activated (or lig-
and-gated) ion channels (1), binding of the lig-
and to the receptor changes the conformation of
the receptor protein. This causes an ion-specific
channel in the receptor protein to open. The re-
sulting flow of ions changes the electric charge
of the cell membrane. Receptors with ligand-
274
Genetic Defects in Ion Channels
More than 20 different disorders due to defec-
tive ion channel proteins resulting from gene
mutations are known. Such disorders include
cystic fibrosis (see p. 276), the long-QT syn-
drome, a special type of deafness, hereditary
hypertension (Liddle syndrome), familial per-
sistant hyperinsulinemic hypoglycemia of in-
fancy, some hereditary muscle diseases, and
malignant hyperthermia (see p. 372), among
other disorders.
A. Long-QT syndrome, a genetic
cardiac arrhythmia
Congenital long-QT syndrome is characterized
by a prolonged QT interval in the electrocardio-
gram (more than 460 ms, corrected for heart
rate), sudden attacks of missed heart beats
(syncopes) or series of rapid heart beats (tor-
sades de pointes), and an increased risk for sud-
den death from ventricular fibrillation in
children and young adults.
B. Different molecular types of
long-QT syndrome
Prolongation of the QT interval in the electro-
cardiogram results from an increase in the du-
ration of the cardiac action potential (1). The
normal potential lasts about 300 ms (phases 1
and 2). The resting membrane potential (phase
3) is reached by progressive inactivation of cal-
cium currents and increasing depletion of
potassium currents, which repolarize the cell.
In phase 0 the cell is quickly depolarized by acti-
vated sodium currents following an excitatory
stimulus.
LQT1 accounts for about half of the patients
with long-QT syndrome. The gene for LQT2 en-
codes a 1195-amino-acid transmembrane pro-
tein responsible for the other major potassium
channel that participates in phase 3 repolariza-
tion (HERG stands for (human-ether-r-go-go-
related gene, a Drosophila homologue). LQT3, a
sodium channel protein, consists of four sub-
units, each containing six transmembrane
domains and a number of phosphate-binding
sites. Homozygosity for LQT1 (KVLQT1 gene) or
LQT5 (KCNE1 gene) causes a form of long-QT
syndrome associated with deafness, the Jervell
and Lange-Nielsen syndrome. (Figure adapted
from Ackerman and Clapham, 1997.) It is impor-
tant to distinguish the different types because
the choice of medication differs.
References
Ackerman, M.J., Clapham, D.D.: Ion channels—
Basic science and clinical disease. New Eng.
J. Med. 336 : 1575 – 1586, 1997.
Keating, M.T., Sanguinetti, M.C.: Molecular
genetic insights into cardiovascular disease.
Science 272 :681 – 685, 1996.
Schulze-Bahr, E., et al.: KCNE1 mutations cause
Jervell and Lange-Nielsen syndrome. Nature
Genet. 17 :267 – 268, 1997.
Schulze-Bahr, E., et al.: The long-QT syndrome.
Current status of molecular mechanisms. Z.
Kardiol. 88 :245 – 254, 1999.
Viskin, S. : Long QT syndromes and torsades de
pointes. Lancet 354 :1625 – 1633, 1999.
Examples of Diseases due to Genetic Ion Channel Defects
Disease Inheritance Ion Channel Gene
Locus
Cystic fibrosis AR CFTR (chloride) 7q31
Long-QT syndrome (6 types) AD KVLQT1 (cardiac potassium) 11p15.5
HERG 7q35 – 36
SCNA5 (cardiac sodium) 3p21
Three other types (see text)
Malignant hyperthermia
Let A and B be p × q and q × r-dimensional matrices, respectively. The size of the
problem depends on the three dimensions p, q, and r. A trivial base case occurs
when p = q = r = 1, where the result is a simple scalar number. Additionally, some
implementations may require considering situations where a dimension is 0. In those
cases the output should be an empty matrix, as will be addressed shortly.
One fairly straightforward way to decompose the problem consists of partitioning
each matrix into four block matrices (forming a 2 × 2 array of block matrices). In
that case their product can be defined as follows:
AB=[ A1,1 A1,2 ] [ B1,1 B1,2 ] A2,1 A2,2 B2,1 B2,2
(6.5)
Notice that the formula is analogous to multiplying two 2 × 2 matrices. For
example, the top-left block of the result (A1,1B1,1 +A1,2B2,1) can be viewed as the
product between the first (block) row of A and the first (block) column of B.
= [ A1,1B1,1 + A1,2B2,1 A1,1B1,2 + A1,2B2,2 ] . A2,1B1,1 + A2,2B2,1 A2,1B1,2 +
A2,2B2,2
The decomposition involves computing eight simpler matrix products. Thus, the
method will invoke itself eight times in the recursive case. The results of each
product need to be added and stacked appropriately in order to form the output
matrix. Listing 6.9 shows a possible implementation. The recursive case first
defines each of the smaller block matrices, adds the simpler products, and builds
the output matrix through the methods vstack and hstack. One of the base cases com-
putes a simple product when p = q = r = 1. In addition, the code also considers the
possibility of receiving empty input matrices, since they appear when partitioning
the matrices in the recursive case if one of the dimensions is equal to one
(obviously, a vector cannot be partitioned into four vectors as described in
(6.5)). Thus, if any of the dimensions is 0 a special base case returns an empty
matrix of dimensions p × r, which can be handled appropriately in Python.
The previous method creates 1 × 1 matrices (in a base case) and progressively
stacks them together to form the final p × r matrix. In addition, note that the
dimensions of the input matrices to the methods are not fixed.
Another more efficient alternative consists of passing the entire matrices A and
B in each call, and specifying the blocks that need to be
188 Introduction to Recursive Programming
Listing 6.9 Divide and conquer matrix multiplication.
1 import numpy as np 2
3
4 def
matrix_mult(A, p = A.shape[0] q = A.shape[1] r = B.shape[1]
ifp==0orq
return np.zeros((p, r))
B):
5 6 7 8 9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
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30
31
32
33 A =
34 B =
35 print(matrix_mult(A, B))
multiplied through appropriate limits passed as parameters, similarly to Listing
1.6. In addition, the result can be stored in a p×r matrix parameter C (passed by
reference). Listing 6.10 shows a possible implementation of this alternative
solution. The method matrix_mult_limits always passes the entire matrices A and B
in each call, storing the result in a p × r matrix (its third parameter).
Additionally, it specifies the submatrices that it will actually multiply through
the rest of the param-
== 0 or r == 0:
elifp==1andq==1andr== 1:
return np.matrix([[A[0, 0] * B[0, 0]]])
else: A11=A[0:p//2,0:q//2] A21=A[p//2:p,0:q//2] A12=A[0:p//2,q//2:q]
A22=A[p//2:p,q//2:q]
B11=B[0:q//2,0:r//2] B21=B[q//2:q,0:r//2] B12=B[0:q//2,r//2:r]
B22=B[q//2:q,r//2:r]
C11 = matrix_mult(A11, B11) C12 = matrix_mult(A11, B12) C21 =
matrix_mult(A21, B11) C22 = matrix_mult(A21, B12)
+ matrix_mult(A12, B21) + matrix_mult(A12, B22) + matrix_mult(A22, B21) +
matrix_mult(A22, B22)
return np.vstack([np.hstack([C11, C12]), np.hstack([C21, C22])])
np.matrix([[2, 3, 1, -3], [4, -2, 1, 2]])
np.matrix([[2, 3, 1], [4, -1, -5], [0, -6, 3], [1, -1, 1]])
Multiple Recursion I: Divide and Conquer 189 Listing 6.10 Alternative divide
and conquer matrix multiplication.
1 import numpy as np
2
3
4 def add_matrices_limits(A, B, C, lp, up, lr, ur):
5 6 7 8 9
10 def 11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43 def 44
45 46 47
for i in range(lp, up + 1): for k in range(lr, ur + C[i,k]=A[i,k]+
matrix_mult_limits(A, B, C, mp = (lp + up) // 2
mq = (lq + uq) // 2
mr = (lr + ur) // 2
1): B[i, k]
lp, up, lq, uq, lr, ur):
if lp == up and lq == uq and lr == ur: C[mp, mr] =
A[mp, mq] * B[mq, mr]
elif lp <= up and lq <= uq and lr <= ur:
M1 = np.zeros((A.shape[0], B.shape[1]))
M2 = np.zeros((A.shape[0], B.shape[1]))
matrix_mult_limits(A, B, M1, lp, mp, lq, mq, lr, mr) matrix_mult_limits(A, B, M2,
lp, mp, mq + 1, uq, lr, mr) add_matrices_limits(M1, M2, C, lp, mp, lr, mr)
matrix_mult_limits(A, B, M1, lp, mp, lq, mq, mr + 1, ur) matrix_mult_limits(
A, B, M2, lp, mp, mq + 1, uq, mr + 1, ur) add_matrices_limits(M1, M2, C, lp, mp, mr
+ 1, ur)
matrix_mult_limits(A, B, M1, mp + 1, up, lq, mq, lr, mr) matrix_mult_limits(
A, B, M2, mp + 1, up, mq + 1, uq, lr, mr) add_matrices_limits(M1, M2, C, mp + 1,
up, lr, mr)
matrix_mult_limits(
A, B, M1, mp + 1, up, lq, mq, mr + 1, ur)
matrix_mult_limits(
A, B, M2, mp + 1, up, mq + 1, uq, mr + 1, ur)
add_matrices_limits(M1, M2, C, mp + 1, up, mr + 1, ur)
matrix_mult_limits_wrapper(A, B):
C = np.zeros((A.shape[0], B.shape[1])) matrix_mult_limits(A, B, C, 0, A.shape[0] -
1,
return C
0, A.shape[1] - 1, 0, B.shape[1] - 1)
190 Introduction to Recursive Programming

eters, which indicate lower and upper limits related to the dimensions p, q, and r.
The base case of matrix_mult_limits occurs when both of the submatrices correspond
to scalar numbers, say ai,j and bj,k. In that case the method simply stores their
product in row i and column k of C. Lastly, the method is not a function, since it
does not return a matrix. Instead, it is a procedure that modifies the parameter C,
where it stores the result. Finally, the iterative method add_matrices_limits adds
the elements of submatrices passed as the first two matrix input parameters, and
stores the result in its third parameter (the submatrices are specified through
parameter limits).
6.5.2 Strassen’s matrix multiplication algorithm
The most expensive arithmetic operation carried out by the previous algorithms is
the scalar multiplication in the base cases. In particular, both require pqr of
these multiplications, similarly to the straightforward iterative version that uses
three loops. It is nevertheless interesting to analyze the time complexity assuming
that the input matrices are n × n. In that case the runtime can be specified
through the following function:
T (n) = ⎧⎪⎨⎪⎩1 if n ≤ 1, (6.6) 8T (n/2) + 4Θ(n2) if n > 1,
since the methods invoke themselves eight times, and need to perform four matrix
additions whose cost is quadratic with respect to n. There- fore, according to the
master theorem (see (3.28)), T(n) ∈ Θ(nlog2 8) = Θ(n3). We will now describe
Strassen’s algorithm, which is a well-known method that can reduce the time
complexity to Θ(nlog2 7) = Θ(n2.807...).
The method also decomposes each of the input matrices into four block matrices as
in the standard algorithm. Thus, AB = C can be expressed as:
[ A1,1 A1,2 ] [ B1,1 B1,2 ]=[ C1,1 C1,2 ] A2,1 A2,2 B2,1 B2,2 C2,1 C2,2
Multiple Recursion I: Divide and Conquer 191 The key to the method is the
definition of the following new matrices
that involve one matrix multiplication operation:
block matrices:
C1,1 =M1 +M4 −M5 +M7, C1,2 = M3 + M5,
C2,1 = M2 + M4,
C2,2 =M1 −M2 +M3 +M6.
M1 = (A1,1 + A2,2)(B1,1 + B2,2), M2 = (A2,1 + A2,2)B1,1,
M3 = A1,1(B1,2 − B2,2), M4 = A2,2(B2,1 − B1,1),
(6.7)
Finally, these matrices can be combined as follows to form the output’s
M5 = (A1,1 + A1,2)B2,2,
M6 = (A2,1 − A1,1)(B1,1 + B1,2), M7 = (A1,2 − A2,2)(B2,1 + B2,2).
The algorithm therefore computes seven products and 18 additions (or subtractions)
in every recursive call. Thus, its runtime cost is described
through:
T(n) = ⎧⎪⎨⎪1 if n ≤ 1, (6.9) ⎪⎩7T (n/2) + 18Θ(n2) if n > 1,
which implies that T(n) = Θ(nlog2 7) = Θ(n2.807...). The algorithm can be faster
than the standard method that runs in Θ(n3) for large values of n. Nevertheless,
for small or medium-sized matrices it may be slower due to the larger
multiplicative constants that play a role in practice.
Finally, from a theoretical point of view, the inputs for this algorithm need to
be square n × n matrices, where n is a power of two. Nev- ertheless, in practice
efficient implementations split the matrices into numerous square submatrices, and
apply the algorithm repeatedly. A simpler, but slower, alternative consists of
padding (i.e., extending) the input matrices with zeros, in order for them to be 2k
× 2k matrices (see Exercise 6.6)
AD
AD, autosomal dominant; AR, autosomal recessive.
(Ryanodine, muscle calcium) 19q13.1
(skeletal-muscle sodium) 17q23 – 25
(About 18 others not listed, adapted from Acker-
man and Clapham, 1997.)
Romano Ward syndrome
(Long-QT syndrome)
Prolonged QT interval in
the electrocardiogram
Syncope
Prolonged QT
Sudden death
Autosomal dominant
Six genes involved
(LQT1 - LQT6)
Torsade de pointes
Type Locus Gene
LQT1 11p15.5 KCNQ1 (KVLQT1)
LQT2 7q35-36 HERG
LQT3 3p21-24 SCNA5
LQT4 4q25-27 unknown
LQT5 21q22.1 KCNE1
LQT6 21q21.1 KCNE2
Long-QT and Deafness (Jervell and
Lange-Nielsen) due to allelic mutations
at LQT1 and LQT5
(autosomal recessive)
1. Main features
2. Electrocardiogram
3. Genetics
A. Long-QT syndrome, a genetic cardiac arrhythmia
+47mV
1
Prolonged cardiac
action potential
2
3
0
Current
clamp
4
–85mV
Normal
0
100
200
300
400
500
Milliseconds
1. Increased duration of cardiac action potential
LQT3 (3q21-24)
SCN5A=Na
II
III
LQT1 (11p15.5)
"
IV
Cell membrane
C
581
N
1
!KPQ
P
N
1
KvLQT1=IKs
C
2016
2. Voltage-activated K-channel delayed
in phase 3
"
LQT2 (7q35-36)
P
P
HERG=IKr
P
P
4. Na-channel fails to inactivate completely
during phase 0
B. Different molecular types of long-QT syndrome
N
1
C
1159
3. Voltage-gated K-channel delayed
in phase 3
Chloride Channel Defects:
Cystic Fibrosis
Cystic fibrosis (mucoviscidosis) is a highly vari-
able multisystemic disorder due to mutations of
the cystic fibrosis transmembrane conduction
regulator gene (CFTR). Cystic fibrosis (CF) is one
of the most frequent autosomal recessive
hereditary diseases in populations of European
origin (about 1 in 2500 newborns). The high
frequency of heterozygotes (1 : 25) is thought to
result from their selective advantage due to re-
duced liability to epidemic diarrhea (cholera).
A. Cystic fibrosis: clinical aspects
The disease primarily affects the bronchial sys-
tem and the gastrointestinal tract. Viscous
mucus formation leading to frequent, recurrent
bronchopulmonic infections and eventually
chronic oxygen deficiency characterize the
common, severe form of the disease. The aver-
age life expectancy in typical CF is about 30
years. The disease may take a less severe, almost
mild course. Congenital bilateral absence of the
vas deferens (CBAVD) occurs in 95% of patients
with CF. It may be the only manifestation in in-
dividuals with different mutant allelic combi-
nations at the CF locus.
B. Positional cloning of the gene for
cystic fibrosis (CF)
The CFTR gene was isolated on the basis of its
chromosomal location (positional cloning) on
the long arm of chromosome 7 at q31 (7q31).
Since the gene could be mapped to the long arm
of chromosome 7 near a marker locus D7S15, a
long-range restriction map comprising about
1500 kb containing the presumptive CF locus
flanked by two marker loci, MET and D7S8, was
constructed. From there a region of 250 kb was
isolated by a combination of chromosome
walking and chromosome jumping. Several
genes were located in this region (candidate
genes) between the marker loci D7S340 and
D7S424. The gene sought was identified by the
finding of mutations of this gene in patients but
not in controls, by comparing with similar
genes in other organisms (evolutionary conser-
vation), by determining its exon/intron struc-
ture, by sequencing it, and by determining the
expression pattern of the gene in different tis-
sues.
C. The CFTR gene and its protein
The CFTR gene is large, extending over 250 kb of
genomic DNA, and is organized into 27 exons
(24 are shown in the diagram) encoding a 6.5 kb
transcript with several alternatively spliced
forms of mRNA. The CFTR protein has 1480
amino acids. It is a membrane-bound chloride
ion channel regulator with several functional
domains: two nucleotide-binding domains (en-
coded by exons 9 – 12 and 19 – 23), a regulatory
domain (exons 12 – 14a), and two transmem-
brane-spanning domains (exons 3 – 7 and 14b –
18). Each of the two transmembrane regions
consists of six transmembrane segments. The
nucleotide-binding domain 1 (NBD1) confers
cAMP-regulated chloride channel activity. The
most common mutation (occurring in 66% of
patients), a deletion of a phenylalanine codon in
position 508 ( ∆ F508), is located here. The pro-
tein is a member of the ATP-binding cassette
(ABC) family of transporters. The R domain con-
tains putative sites for protein kinase A and pro-
tein kinase C phosphorylation. CFTR is widely
expressed in epithelial cells. The more than 800
different mutations observed in the CFTR gene
(see http://www.genet.sickkids.on.ca/cftr) can be
grouped into five different functional classes:
(i) abolished synthesis of full-length protein, (ii)
block in protein processing, (iii) reduced
chloride channel regulation, (iv) reduced
chloride channel conductance, (v) reduced
amount of normal CFTR protein. The underlying
genetic defects include missense mutations,
nonsense mutations, RNA splicing mutations,
and deletions. Aside from ∆ F508 in about 66%
of patients, the most frequent mutations world-
wide are G542X (2.4%), G551D (1.6%), N1303K
(1.3%), and W1282X (1.2%).
References
Chillon, M., et al.: Mutations in the cystic fibro-
sis gene in patients with congenital absence
of the vas deferens. N. Engl. J. Med.
332 :1475 – 1480, 1995.
Rosenbluth, D.B., Brody, S. L.: Cystic fibrosis, pp.
329 – 338, In: J.L. Jameson, ed., Principles of
Molecular Medicine. Human Press, Totowa,
N.J., 1998.
Rosenstein, B.J., Zeitline, P.C.: Cystic fibrosis.
Lancet 351 :277 – 282, 1998.
Tsui, L.C.: The spectrum of cystic fibrosis muta-
tions. Trends Genet. 8 :392 – 398, 1992.
All rights reserved. Usage subject to terms and conditions of license.Chloride
Channel Defects: Cystic Fibrosis
277278
Rhodopsin, a Photoreceptor
The human retina contains about 110 million
rod cells for vision in the dark and 6 million
cone cells for color vision in the light. These
cells contain photoreceptors that convert light
into a nerve impulse. Rhodopsin is the photore-
ceptor for weak light. The light-transmitting
system consists of numerous components
coded for by genes that are similar in structure
and function to genes for other transmembrane
signal-transmitting molecules.
A. Rod cells
A rod cell consists of an outer segment with a
photoreceptor region and an inner segment
comprising cell nucleus and cytoplasm with en-
doplasmic reticulum, Golgi apparatus, and mi-
tochondria. The outer segment contains about
1000 disks with rhodopsin molecules in the
membrane. In the periphery, the approximately
16 nm thick disks are folded by the protein pe-
ripherin. (Diagram after Stryer, 1995).
B. Photoactivation
In 1958, George Wald and co-workers dis-
covered that light isomerizes 11-cis-retinal (1)
very rapidly into all-trans-retinal (2), a form
that practically does not exist in the dark
(!1 molecule/1000 years). The light-induced
structural change is so great that the resulting
atomic motion can trigger a reliable and repro-
ducible nerve impulse. The absorption spec-
trum of rhodopsin (3) corresponds to the spec-
trum of sunlight, with an optimum at a
wavelength of 500 nm. Although vertebrates,
arthropods, and mollusks have anatomically
quite different types of eyes, all three phyla use
11-cis-retinal for photoactivation.
D. Rhodopsin
Rhodopsin is a seven-helix transmembrane
protein with binding sites for functionally im-
portant molecules such as transducin, rhodop-
sin kinase, and arrestin on the cytosol side. The
binding site of the light-sensitive molecule
(chromophore) is lysine in position 296 of the
seventh transmembrane domain. The light-ab-
sorbing group consists of 11-cis-retinal. The
amino end of rhodopsin is located in the disk in-
terspaces, and the carboxy end on the cytosol
side. About half of the molecule is contained in
the seven transmembrane hydrophobic do-
mains, one-fourth in the disk interspaces and
one-fourth on the cytosol side.
E. cGMP as transmitter in the
vizualization process
The light cascade ends with rapid hydrolysis of
cGMP, the internal transmitter in visualization.
This leads to rapid closure of the sodium ion
channels and hyperpolarization of the mem-
brane to initiate nerve impulse, which is trans-
mitted as a signal to the brain.
References
Palczewski, K. et al.: Crystal structure of
rhodopsin: A G protein-coupled receptor.
Science 289 :739 –745, 2000.
Schoenlein, R.W., et al.: The first step in vision:
Femtosecond isomerization of rhodopsin.
Science 254 :412 – 415, 1991.
Stryer, L.: Biochemistry. 4 th ed. W.H. Freeman,
New York, 1995.
Stryer, L.: Molecular basis of visual excitation.
Cold Spring Harbor Symp Quant Biol. 53 :28 –
294, 1988.
C. Light cascade
Photoactivated rhodopsin triggers a series of
enzymatic steps (light cascade). First, a signal-
transmitting protein of visualization, trans-
ducin, is activated by photoactivated rhodopsin.
Transducin belongs to the G protein family, i.e.,
it can assume an inactive GDP and an active GTP
form. GTP activates phosphodiesterase. This
very rapidly hydrolyzes cGMP and lowers the
cGMP concentration in cytosol, which leads to
closure of the sodium ion channels. Immedi-
ately thereafter, phosphodiesterase is inacti-
vated by means of a G protein cycle.
All rights reserved. Usage subject to terms and conditions of license.Rhodopsin, a
Photoreceptor
279280
Mutations in Rhodopsin
Retinitis pigmentosa (RP) is a genetically heter-
ogeneous group of diseases that lead to
pigmental degeneration of the retina and pro-
gressive blindness. Numerous mutations in the
rhodopsin gene have been shown to be the
cause of different forms of RP. Mutations in
other genes coding for proteins of the light cas-
cade may also cause retinitis pigmentosa.
A. Retinitis pigmentosa
The fundus of the eye shows distinct displace-
ment of pigmentation, with irregular hyper-
pigmentation and depigmentation. The papilla
(optic disk) shows waxy yellow discoloration.
The loss of vision, especially in dim light (night
blindness), proceeds from the periphery to the
center at different rates depending on the form
of the disease, until only a very narrow central
visual field remains. (Photograph from E. Zren-
ner, Tübingen.)
B. Point mutation in codon 23
The first point mutation demonstrated in the
rhodopsin gene (Dryja et al., 1990) was a trans-
version from cytosine to adenine in codon
23. This changed the codon CCC for proline (Pro)
into CAC for histidine (His). Since the proline in
position 23 occurs in more than ten related G
protein receptors, it must be very important for
normal function.
C. Mutations in rhodopsin
The gene locus for rhodopsin (RHO) in man lies
on the long arm of chromosome 3 in region 2,
band 1.4 (3q21.4). Dominant and autosomal re-
cessive inherited mutations have been demon-
strated in humans. Most mutations lead to the
exchange of an amino acid, although deletions
may also occur. Of the 348 amino acids of
rhodopsin, 38 are identical (invariant) at
various positions in vertebrates. More than 100
different mutations are known for autosomal
dominant inherited RP. An increasing number
of mutations are recognized to cause autosomal
recessive RP. In addition, mutations in several
other gene loci have been recognized to lead to
retinitis pigmentosa, e. g. mutations in the gene
for peripherin on the short arm of chromosome
6 in humans (6p) and a locus in the centromeric
region of chromosome 8. Other photoreceptor
gene disease loci are the α and β subunits of
phosphodiesterase (PDE).
D. Demonstration of a mutation in
codon 23 by means of
oligonucleotides after PCR
This pedigree (1) with autosomal dominant in-
herited retinitis pigmentosa due to mutation in
codon 23 includes 13 affected individuals in
three generations (affected females, black
circles; affected males, black squares). Using
polymerase chain reaction (PCR) (see p. 166),
Dryja et al. (1990) demonstrated the mutation
in amplified fragments of exon 1 (2). The nor-
mal oligonucleotide corresponds to the normal
sequence between codons 26 and 20. The mu-
tant sequence of the oligomere RP contains the
mutant sequence CAC. All affected individuals
gave a hybridization signal with the RP oli-
gomer (2) (II-2, II-12, and III-4 were not ex-
amined), whereas unaffected individuals did
not (see p. 408 for demonstration of a point mu-
tation with oligonucleotides).
References
Barkur, S. S. : Retinitis pigmentosa and related
disorders. Am J Med Genet. 52 :467 – 474,
1994.
Dryja, T.P.: Retinitis pigmentosa, pp. 4297 –
4309, In: C.R. Scriver, et al., eds., The Meta-
bolic and Molecular Bases of Inherited Dis-
ease. 7 th ed. McGraw-Hill, New York, 1995.
Dryja, T.P., et al.: A point mutation of the
rhodopsin gene in one form of retinitis pig-
mentosa. Nature 343 :364 – 366, 1990.
McInnes, R.R., Bascom, R.A.: Retinal genetics: a
nullifying effect for rhodopsin. Nature
Genetics 1 :155 – 157, 1992.
Wright, A.F.: New insights into genetic eye dis-
ease. Trends Genet. 8 :85 – 91, 1992.
All rights reserved. Usage subject to terms and conditions of license.Mutations in
Rhodopsin
281282
Color Vision
As suggested by Thomas Young in 1802, color
vision in humans is mediated by three receptor
types in the cone cells of the retina, one each for
blue, green, and red.
A. Genes for photoreceptor proteins in
cones
The gene for the blue receptor is autosomal; the
genes for the red and green receptors are X
chromosomal. The absorption spectra of the
three receptors show maxima of 426 nm for
blue, about 530 for green, and about 550 for red.
The red receptor was discovered to be polymor-
phic, with two somewhat different absorption
maxima at 552 and 557 nm.
B. Evolution of the genes for visual
pigment photoreceptors
The photoreceptor genes arose from a single an-
cestral gene (protogene). The rhodopsin–trans-
ducin pair is found in invertebrates and is at
least 700 million years old. The blue receptor is
almost as old as rhodopsin, about 500 million
years. The separation into a receptor for green
and one for red must have occurred only about
30 million years ago, after the Old World and
New World apes separated, since man and the
Old World apes have three cone pigments
whereas New World apes have two.
C. Structural similarity of the visual
pigments
In 1986, J. Nathans and co-workers sequenced
the genes for color photoreceptors and ob-
served marked structural similarities, es-
pecially of the green and red receptor genes.
Here the gene products (the receptors) are
shown and their similarities are compared. The
dark dots indicate variant amino acids; the light
dots are identical amino acids, given in percent-
ages.
D. Polymorphism in the photoreceptor
for red
A. G. Motulsky and co-workers (Winderickx et
al., 1992) demonstrated variant codons in three
regions of the red receptor gene (1). Serine was
found at position 180 in 60% of the investigated
males; alanine in 40%. Position 230 showed
polymorphism of isoleucine (Ile) and threonine
(Thr); position 233 of alanine (Ala) and serine
(Ser) (2). Differences in red color perception
could be demonstrated by the color-mixing test
procedure of Raleigh (3).
E. Normal and defective red–green
vision
One gene for red and one to three genes for
green lie close together on the long arm of the X
chromosome in humans (1). Since the
sequences of these genes are very similar, un-
equal crossing-over is not infrequent (2). Inter-
genic crossing-over leads to loss (green blind-
ness) or duplication; intragenic crossing-over
leads to a hybrid gene (red blindness). Green
blindness results from loss of a gene for the
green receptor; red blindness, from a defective
or absent red receptor. With red–green blind-
ness, neither a normal red nor a normal green
receptor is present. About 1% of all men are red–
green blind and about 2% are green blind. About
8% show weakness in differentiating red from
green.
References
Kohl, S. , et al.: Total colour blindness is caused
by mutations in the gene encoding the α -
subunit of the cone photoreceptor cGMP-
gated cation channel. Nature Genet. 19 :257 –
259, 1998.
Motulsky, A.G., Deeb, S. S.: Color vision and its
genetic defects, pp. 4275 – 4295. In: C.R.
Scriver, et al., eds., The Metabolic and
Molecular Bases of Inherited Disease. 7 th ed.
McGraw-Hill, New York, 1995.
Nathans, J., Thomas, D., Hogness, D.S. : Molecu-
lar genetics of human color vision: the
genes encoding blue, green, and red pig-
ments. Science 232 :193 – 202, 1986.
Neitz, M., Neitz, J.: Numbers and ratios of visual
pigment genes for normal red-green color
vision. Science 267 :1013 – 1016, 1995.
Winderickx, J., et al.: Polymorphism in red pho-
topigment underlies variation in colour
matching. Nature 356 :431 – 433, 1992.
Wissinger, B., Sharpe, L.T.: New aspects of an old
theme: The genetic basis of human color vi-
sion. Am. J. Hum. Genet. 63 :1257 – 1262,
1998.
Wissinger, B., et al.: Human rod monochro-
macy: linkage analysis and mapping of a
cone photoreceptor expressed candidate
gene on chromosome 2q11. Genomics
51 : 325 – 331, 1998.
All rights reserved. Usage subject to terms and conditions of license.Color Vision
283284
Hearing and Deafness
Acoustic signals are essential for an animal’s
ability to respond appropriately to its environ-
ment. Hearing is orchestrated by a large en-
semble of proteins acting in concert. Special-
ized sensory cells in the cochlea of the inner ear
process the incoming sound waves, converting
them into cellular information that is relayed to
the brain via the acoustic nerve. A missing or
defective protein involved in the hearing
process results in hearing loss. Hearing loss is
common in humans. One out of 1000 newborns
lacks the ability to hear or has severely impaired
hearing. Two categories of genetic hearing loss
can be distinguished: nonsyndromic and syn-
dromic. In the former category the genetic de-
fect is limited to the ear; in the latter the ear is
one of several organ systems affected.
The types of genes implicated when defective as
the cause of nonsyndromic hearing loss include
those encoding proteins involved in cy-
toskeletal structure, transcription factors, ion
channels (potassium channel), and intercellular
gap channels composed of junction connexins.
A. The main components of the ear
The auditory system consists of the outer ear,
the middle ear, and the inner ear. Sound waves
are funneled through the outer ear (auricle) and
transmitted through the external ear canal to
the tympanic membrane, which they cause to
vibrate. These vibrations are transmitted
through the tympanic cavity of the middle ear
by a chain of three movable bones, the malleus,
the incus, and the stapes. Three major cavities
form the inner ear: the vestibule, the cochlea,
and the semicircular canals. The chochlea is the
site where auditory signals are processed. The
cochlea contains a membranous labyrinth filled
with a fluid, the endolymph. The vestibular ap-
paratus includes three semicircular canals
oriented at 90Њ degree angles to each other.
They respond to rotatory and linear accelera-
tion. Signals received here are transmitted via
the vestibular nerve, which fuses with the
cochlear nerve to form the acoustic nerve. The
latter transmits the information to the brain.
B. The cochlea
The cochlea contains the cochlear duct, which
forms the organ of Corti. The organ of Corti con-
verts sound waves in the endolymph of the
cochlea into intracellular signals. These are
transmitted to auditory regions of the brain.
The organ of Corti contains two types of sensory
cells: one row of inner hair cells and three rows
of outer hair cells. The inner hair cells are pure
receptor cells. Vibrations induced by sound lead
to slight deflections of the stereocilia and open
potassium channels at the tips of the stereocilia.
The influx of potassium ions at the tips of the
cilia of the hair cells (see C) causes a change in
membrane potential that results in a nerve im-
pulse, which is transmitted as an auditory sig-
nal to the auditory cortex of the brain.
Potassium ions are recycled to the supporting
cells and the spiral ligament into the en-
dolymph of the scala media. The tectorial mem-
brane amplifies the sound waves as a resonator.
(Figure adapted and redrawn from Willems,
2000.)
C. The outer hair cell
The outer hair cells combine sensory function
with the ability to elongate and contract when
acoustically stimulated. The apical pole of a hair
cell carries an array of about 100 cylindrical
stereocilia in a V-shaped arrangement. Each
stereocilium contains an actin molecule, which
enables it to elongate or to contract. The tips of
the stereocilia are connected by tip links. The
potassium channels are formed by the KCNQ4
protein (yellow) and by connexins (red). Impor-
tant for the structural integrity and dynamics of
the hair cells is a cytoskeleton involving actin,
myosin 7A, myosin 15, and the protein di-
aphanous. (Figure adapted and redrawn from
Willems, 2000.)
D. Chromosomal locations of human
deafness genes
Almost every human chromosome harbors at
least one gene involved in nonsyndromic
monogenic hearing loss. The diagrammatic
presentation shown here is limited to nonsyn-
dromic hearing loss.
References
Robertson N.D., Morton, R.N.D.: Beginning of a
molecular era in hearing and deafness. Clin.
Genet. 55 :149 – 159, 1999.
Steel, K.P., Bussoli, T.J.: Deafness genes. Expres-
sions of surprise. Trends Genet. 15 :207 –
211, 1999.
variază la diferite grupe de insecte, putând ajunge la unele specii până la 50 %

(la
larvele unor specii de chironomide).
C i r c u l a ţ i a s â n g e l u i la insecte se face cu ajutorul cămăruţelor
inimii şi
diafragmelor dorsală şi ventrală. Sub acţiunea muşchilor cămăruţele inimii se
dilată şi se
contractă consecutiv, iar sângele este împins dintr-o cămăruţă în alta până la
aortă. În
momentul dilatării (diastola), toate valvulele unei cămăruţe se deschid şi sângele
pătrunde în interior, atât din sinusul pericardial cât şi din cămăruţă posterioară.
În
momentul contractării (sistola) este deschisă numai valvula din faţă, prin care
sângele
este împins în cămăruţa anterioară, iar celelalte valvule sunt închise, oprind
refluxul
(fig. 24). Frecvenţa contracţiilor cămăruţelor inimii variază între 30 - 140 pe
minut, în
funcţie de specia de insectă. Sângele ajuns în aortă se revarsă prin orificiul ei
anterior în
cavitatea cefalică şi de acolo în tot corpul. Ridicarea sângelui din sinusul
perineural în
sinusul visceral, iar din acesta în cel pericardial se realizează prin contractarea
ritmică a
diafragmelor ventrală şi dorsală, care prezintă lateral şi anterior spaţii
deschise, prin care
sinusurile comunică între ele. Sângele ajuns în sinusul pericardial pătrunde din
nou în
vasul dorsal, circulând dinapoi-înainte şi de jos în sus. Organele pulsatorii ajută
în
circulaţia sângelui, împingându-l în antene, picioare şi aripi.
Fig. 24 - Schema funcţionării inimii la insecte :
a - camera inimii în stare de sistolă; b - camera inimii în stare
de diastolă; Os - ostiole; Va - valvule
(după P o s p e l o v).
4.6. SISTEMUL RESPIRATOR
La marea majoritate a insectelor, la fel ca la myriapode, respiraţia este t r a-
h e a n ă. Ea se realizează prin intermediul unui sistem de tuburi ramificate
numite
t r a h e i, care comunică cu exteriorul prin nişte orificii denumite s t i g m e
(fig. 25).
Sistemul trahean este caracteristic insectelor terestre; la insectele acvatice (la
larvele de
ephemeroptere, plecoptere, odonate etc.) respiraţia se face prin branhii traheene.
S t i g m e l e sau s p i r a c o l e l e t r a h e e n e (stigma, spiracula) sunt
orificii ale traheelor, situate pe părţile laterale ale corpului, câte o pereche pe
fiecare
segment, lipsind, în general, pe cap, protorace şi segmentul al 9-lea abdominal.
Numărul
29stigmelor variază la diferite grupe de insecte (1 - 2 perechi stigme toracice şi
2 perechi
pe segmentele abdominale la thysanoptere, o pereche de stigme toracice şi 6 perechi
abdominale la anoplure etc.).
Fig. 25 - Sistemul trahean la insecte (Periplaneta):
a - văzut ventral; b - văzut dorsal; Tr - trahei; St - stigme
(după M i a l l şi D e n n y).
Stigmele sunt mărginite de un cadru chitinos, îngroşat, numit p e r i t r e m, iar
orificiul lor continuă cu o invaginare sub formă de cameră, denumită a t r i u m
(fig.
26). Pereţii interni ai camerei atriale sunt căptuşiţi cu perişori sau excrescenţe
chitinoase, care au rol de filtru. Atriul uneori este format din două camere.
Deschiderea
şi închiderea stigmelor se face prin intermediul unor dispozitive speciale,
acţionate de
muşchii închizători şi deschizători.
Fig. 26 - Secţiune longitudinală printr-o stigmă abdominală la furnică (Formica) :
a - stigmă deschisă; b - stigmă închisă; Ds - deschiderea stigmei; Ca - camera
anterioară; Cp - camera posterioară; Mi - muşchi închizător; Md - muşchi
deschizător; Tr - trahee (după J a n e t).
30T r a h e i l e sunt de origine ectodermică şi reprezintă invaginări ale
tegumentului. Peretele traheii este alcătuit din m e m b r a n a b a z a l ă, un
strat de
celule hipodermice, numit e p i t e l i u sau m a t r i c e a şi i n t i m a, care
corespunde cuticulei tegumentului. Cel mai dezvoltat înveliş cuticular este
exocuticula,
care este îngroşată sub formă de spirală şi poartă denumirea de t e n i-
d i e (tenidium). Epicuticula şi endocuticula, mai dezvoltate în tuburile traheene
groase, sunt subţiate mult în traheele subţiri. Tenidia are rolul de a menţine
tubul
trahean deschis, de a împiedica turtirea lui.
De la camera atrială porneşte o trahee scurtă, care se ramifică în 3 ramuri : o
r a m u r ă d o r s a l ă, care deserveşte inima, musculatura şi tegumentul; o r a
m u- r ă
v e n t r a l ă, care merge la sitemul nervos, musculatură şi tegument şi o r a m
u- r ă v
i s c e r a l ă spre intestin şi celelalte organe viscerale. Ramurile traheene, în
continuare, se divid dicotomic în trahei din ce în ce mai fine şi înguste, până la
dimensiunile capilarelor, cu un diametru mai mic de un micron, numite t r a h e o l
e
(fig. 27). Acestea din urmă sunt lipsite de tenidie şi pătrund în interiorul
celulelor
corpului, având pereţii permeabili pentru gaze şi lichide.
La numeroase insecte (unele hymenoptere, diptere, orthoptere etc.) tuburile
traheene longitudinale prezintă dilatări, mai ales în regiunea abdomenului, numite
s a c i a e r i e n i, care servesc ca rezervoare de aer în timpul zborului.
Fig. 27 - Schema traheolelor:
Fm - fibră musculară; Te - traheole;
Tr - trahee (după K e l l e r).
F i z i o l o g i a r e s p i r a ţ i e i. Procesul respiraţiei la insecte se
realizează
prin aerisirea traheelor, respectiv pătrunderea aerului în ţesuturi şi eliminarea
bioxidului
de carbon. Expiraţia este activă, efectuându-se prin apăsarea sistemului trahean
datorită
contractării muşchilor dorso-ventrali, care apropie tergitele de sternite, şi prin
contractarea muşchilor longitudinali, care scurtează abdomenul. Inspiraţia,
datorită
revenirii abdomenului la normal, este pasivă. La inspiraţie, aerul pătrunde prin
stigme în
trahei şi traheole. La nivelul acestora din urmă se realizează difuziunea
oxigenului în
celule.
La unele insecte (apterygote) toate stigmele participă atât la inspiraţie cât şi la
expiraţie. La altele (Acrididae), unele stigme servesc la inspiraţie, iar altele la
expiraţie,
închizându-se ritmic. Frecvenţa mişcărilor respiratorii la insecte variază între 12
- 100
pe minut, în funcţie de specie, factorii mediului ambiant (temperatură, umiditate
etc.) şi
intensitatea activităţii (repaus, mişcare).
În timpul respiraţiei se produce oxidarea substanţelor proteice, a grăsimilor şi a
hidraţilor de carbon din ţesuturi şi eliberarea bioxidului de carbon. Reacţia
oxidării este
exotermică şi exodinamică, eliberându-se energia şi căldura necesare întreţinerii
31proceselor vitale. Eliminarea bioxidului de carbon se face prin trahei şi numai
în mică
parte (până la 25 %) este difuzat prin tegument în mediul exterior.
La unele insecte inferioare (apterygote), ca şi la unele larve endoparazite (unele
hymenoptere şi diptere), respiraţia este cutanee, oxigenul difuzând prin tegument.
4.7.
SISTEMUL EXCRETOR
În urma proceselor metabolice care au loc în ţesuturi şi organe, în afara
sistemului digestiv, rezultă unele substanţe inerte sau vătămătoare organismului
(acizi
urici, oxalaţi, carbonaţi etc.), care sunt eliminate sub formă de e x c r e t e
prin sistemul
excretor. Funcţia de excreţie, la insecte, este îndeplinită de tuburile lui
Malpighi, corpul
adipos şi alte organe.
T u b u r i l e l u i M a l p i g h i, principalul organ de excreţie, sunt de
origine
ectodermică. Ele se prezintă sub forma unor tuburi lungi şi subţiri, închise la
capetele
libere şi deschise la baza lor de fixare, înapoia valvulei pilorice, aproape de
limita
intestinului mediu cu cel posterior (v. fig. 22). Numărul tuburilor lui Malpighi
variază
mult după grupele de insecte. Astfel, coccidele au 2 tuburi, thysanopterele,
homopterele,
dipterele şi lepidopterele 4, coleopterele 4 - 6, orthopterele 90 - 120 etc.
Din punct
de vedere histologic tuburile lui Malpighi sunt formate dintr-un singur strat de
celule,
mari, epiteliale, învelite de membrana bazală. Tuburile au o musculatură proprie,
care le
asigură mişcările în cavitatea corpului. Din punct de vedere fiziologic, tuburile
lui
Malpighi îndeplinesc rolul de rinichi, extrăgând din sânge acidul uric şi
diferitele săruri.
Aceste produse de excreţie ajung prin canalele tuburilor în intestinul posterior,
de unde
sunt eliminate prin orificiul anal odată cu excrementele. La unele insecte,
tuburile lui
Malpighi îndeplinesc şi alte funcţii ca : organe secretoare de mătase, care servesc
la
confecţionartea coconilor (unele specii de Planipennia, unele coleoptere etc.) sau
organe secretoare a unor fermenţi. La fungivoride (Diptera) partea dorsală a
tuburilor
lui Malpighi este modificată în organe luminoase.
C o r p u l a d i p o s sau c o r p u l g r a s este răspândit în cavitatea
celomică
sub forma a două straturi : unul sub tegument - s t r a t u l p a r i e t a l şi
unul în jurul
tubului digestiv - s t r a t u l p e r i v i s c e r a l. Corpul gras este alcătuit
dintr-o
aglomerare de celule sferice sau poliedrice, cu nucleul mic. Culoarea corpului gras
este
variabilă : albă, verzuie, galbenă, portocalie.
Corpul gras îndeplineşte mai multe funcţii, dintre care cele mai importante sunt
următoarele : ca organ excretor şi ca acumulator de substanţe de rezervă. Astfel,
corpul
gras înmagazinează, ca un rinichi de acumulare, diferite săruri vătămătoare
organismului sub formă de cristale, care pot rămâne în interiorul lui toată viaţa
insectei
sau sunt eliminate prin intermediul tuburilor lui Malpighi. Ca substanţe de rezervă
se
acumulează în corpul gras grăsimile, proteinele şi mai ales glicogenul.
C e l u l e l e p e r i c a r d i a l e sau n e f r o c i t e l e se prezintă sub
forma
unor mase celulare, situate în apropierea vasului sanguin, pe fibrele sau între
fibrele
diafragmei dorsale. Ele extrag din sânge diferite corpuri străine (unele substanţe
proteice, clorofila etc.).
G l a n d e l e l a b i a l e se întâlnesc la unele grupe de insecte inferioare
(collembole - poduride etc.), la care tuburile lui Malpighi lipsesc. Excretele lor
sunt
eliminate printr-un canal care se deschide la baza buzei superioare.
324.8. SISTEMUL SECRETOR
Sistemul secretor al insectelor este alcătuit din două tipuri de glande: exocrine
şi
endocrine. Secreţiile glandelor exocrine ajung la diferite organe prin intermediul
unor
canale speciale. Glandele endocrine sunt lipsite de canale, iar secreţiile lor sunt
transportate cu ajutorul sângelui în toate regiunile corpului.
Glandele exocrine sunt foarte diferite ca origine şi funcţie. Ele sunt repartizate
în diferite regiuni ale corpului insectelor (în tegument, tubul digestiv, organele
genitale
etc.). Unele dintre glandele exocrine sunt prezentate în continuare.
G l a n d e l e c e r i e r e, care secretă ceara, se întâlnesc la unele grupe de
homoptere (coccide, aleurodide, afide).
G l a n d e l e l a c c i p a r e se găsesc la unele specii de coccide (Laccifer
lacca). Ele secretă lacuri speciale, care reprezintă un amestec de răşini şi ceară.
G l a n d e l e r e p u l s i v e secretă substanţe cu mirosuri puternice şi
respingătoare, care au rol de apărare. Ele sunt situate pe torace şi abdomen (la
Eurygaster), la articulaţiile picioarelor (la Meloe, Coccinella) etc.
G l a n d e l e p r o d u c ă t o a r e d e f e r o m o n i, frecvente la numeroase
specii de insecte, sunt formaţiuni glandulare, de o mare varietate, situate în
diferite
regiuni ale corpului (pe aripi, pe abdomen, în abdomen etc.). Secreţiile acestor
glande,
substanţe chimice prin intermediul cărora indivizii aceleeaşi specii comunică între
ei,
sunt cunoscute sub denumirea de f e r o m o n i. Ei sunt în mod tipic odoriferi şi
acţionează direct asupra sistemului nervos al individului receptor.
În funcţie de semnificaţia mesajului, feromonii sunt împărţiţi în două mari grupe:
feromoni de dezvoltare (metabolici) şi feromoni de acţiune (de declanşare).
F e r o m o n i i d e d e z v o l t a r e, care induc la indivizii receptori unele
modificări metabolice sau de dezvoltare.
F e r o m o n i d e a c ţ i u n e, care antrenează schimbări de comportament.
Cele mai importante tipuri sunt redate mai jos.
F e r o m o n i d e b a l i z a j, cu care este marcat intinerariul de deplasare
spre
sursa de hrană etc. (la insectele sociale, la carii de scoarţă şi de lemn).
F e r o m o n i d e o v i p o z i ţ i e, cu care sunt marcate locurile propice
(ţânţari) sau "interzise" (muştele fructelor) pentru depunerea ouălor.
F e r o m o n i d e a l a r m ă, care determină fuga (dispersarea) indivizilor
populaţiei în momentul atacului unui prădător (la afide).
F e r o m o n i d e a g r e g a r e, care asigură concentrarea populaţiei în
vederea migraţiei sau populării unui biotip (lăcuste, carii de lemn şi scoarţă) şi
coeziunea familiei (la insectele sociale).
F e r o m o n i s e x u a l i care mediază relaţiile dintre cele două sexe înainte,
în timpul şi după împerechere. Sunt cunoscute trei grupe de feromoni sexuali.
Atractanţi sexuali, produşi de unul din sexe pentru atragerea sexului opus în
vederea împerecherii. Se cunosc, până în prezent, atractanţi sexuali la numeroase
specii
de insecte din ordinele : Lepidoptera, Coleoptera, Hymenoptera etc. În majoritatea
cazurilor, feromonii sunt produşi de femelă şi au acţiune atractivă asupra
masculului,
dar sunt cazuri când feromonul este produs de mascul şi exercită atracţie asupra
femelei.
La unele specii, reduse ca număr, feromonii sunt produşi şi de mascul şi de femelă.
Atractanţii sexuali au o mare însemnătate practică, utilizându-se îndeosebi în
prognoză şi combaterea unor specii de insecte dăunătoare.
Afrodisiaci, produşi de către masculii atraşi, cu scopul de a excita femelele şi de
a le determina să accepte împerecherea.
33Repelenţi sexuali, produşi în aparatul genital al masculului şi introduşi odată
cu
sperma în cel al femelei. Se marchează astfel femelele fecundate şi îi determină pe
ceilalţi masculi să le evite.
La insecte se întâlnesc şi alte tipuri de glande, dintre care menţionăm: g l a n- d
e
u r t i c a n t e (la larvele unor specii de lepidoptere: Pythiocampa, Lymantria
etc.), cu
rol de apărare: g l a n d e s e r i c i p a r e, din a căror secreţie se formează
firul de
mătase (la unele larve de lepidoptere : Bombyx, Antheraea etc.); g l a n d e
p r o d u c ă t o a r e d e v e n i n (la unele specii de hymenoptere: Apis, Vespa,
Formica), care servesc la apărare etc.
Glandele endocrine. Secreţiile acestor glande, cunoscute sub numele de
h o r m o n i sau i n c r e t e, au un mare rol în viaţa insectelor în ce priveşte
dezvoltarea larvară, năpârlirea, diapauza,
metamorfoza etc. Au fost studiate
următoarele tipuri de glande endocrine : celulele neurosecretorii ale ganglionilor
cerebroizi, glandele protoracale, corpora allata şi corpora cardiaca.
C e l u l e l e n e u r o s e c r e t o r i i a l e g a n g l i o n i l o r c e r e
b r o-
i z i secretă h o r m o n u l c r e i e r u l u i. Rolul acestui hormon se
consideră că este
foarte diferit. În unele cazuri, controlează şi intensifică activitatea glandelor
protoracale; în alte cazuri determină încetarea activităţii acestora din urmă la
larve şi
pupe, frânează creşterea şi dezvoltarea şi determină intrarea şi ieşirea din
diapauză.
Hormonul creierului se poate acumula în corpora cardiaca şi corpora allata,
determinând
activizarea acestora.
G l a n d e l e p r o t o r a c a l e sunt reprezentate printr-o pereche de glande,
situate în partea inferioară a protoracelui, pe laturile ganglionilor respectivi,
cu care sunt
în legătură. Aceste glande secretă h o r m o n u l n ă p â r l i r i i,
m e t a m o r f o z
e i sau e c d y s o n, care determină întreruperea diapauzei şi reglementează
năpârlirile
şi, în general, toată dezvoltarea larvară.
C o r p o r a a l l a t a se prezintă sub forma a doi lobi, situaţi deasupra
tubului
digestiv anterior, în partea posterioară a ganglionilor cerebroizi. Aceste glande,
care se
întâlnesc la larvele şi la adulţii insectelor superioare (Pterygota), precum şi la
unele
insecte inferioare (Apterygota - Diplura), secretă h o r m o n u l j u v e n i l
sau n e o t
e n i n. Acest hormon intervine în morfogeneză şi condiţionează creşterea
proporţională
a larvelor. Reducerea concentraţiei hormonului juvenil în sânge determină la larve
transformarea în pupe şi apoi în insecte adulte.
C o r p o r a c a r d i a c a apare sub forma a doi lobi, situaţi în partea
anterioară
a ganglionilor cerebroizi. Rolul acestor glande este mai puţin cunoscut; se
consideră că
ele reglementează activitatea glandelor protoracale.
Activitatea glandelor endocrine se desfăşoară în complex, sub controlul direct al
sistemului nervos central şi sub influenţa factorilor de mediu.
Cunoaşterea sistemului endocrin şi a fiziologiei sale prezintă o mare valoare
practică. Pe aceasta se bazează combaterea biologică a insectelor cu ajutorul
produselor
hormonale.
4.9. SISTEMUL NERVOS
Sistemul nervos al insectelor, ca şi al celorlalte arthropode, este de tip
scalariform. El este de origine ectodermică şi se compune din sistemul nervos
central,
sistemul nervos simpatic şi sistemul nervos periferic.
S i s t e m u l n e r v o s c e n t r a l este alcătuit dintr-un lanţ dublu de
ganglioni, uniţi între ei longitudinal prin fibre nervoase numite c o n e c t i v e
şi
transversal prin c o m i s u r i.
34Sistemul nervos central este constituit din 3 grupe de ganglioni : ganglionii
cerebroizi, ganglionii subesofagieni şi ganglionii lanţului nervos ventral (fig.
28).
G a n g l i o n i i c e r e b r o i z i sau s u p r a e s o f a g i e n i, care
formează
creierul la insecte, sunt aşezaţi în capsula cefalică, deasupra esofagului. Ei sunt
constituiţi din 3 perechi de ganglioni: protocerebrum, deutocerebrum şi
tritocerebrum.
Protocerebrum, corespunzător primului segment cefalic (acronul), reprezintă
partea cea mai dezvoltată a creierului şi trimite nervi la organele vizuale (ochii
compuşi
şi ocelii).
Deutocerebrum, corespunzător segmentului antenal, inervează antenele, iar
tritocerebrum, corespunzător segmentului labral, trimite nervi la buza superioară
şi
regiunea frontală.
Prin inelul periesofagian creierul este legat de ganglionii subesofagieni.
G a n g l i o n i i s u b e s o f a g i e n i sunt situaţi sub esofag şi s-au
format din
contopirea a trei perechi de ganglioni, care corespund segmentelor : mandibular,
maxilar şi labial. Ei inervează mandibulele, maxilele şi buza inferioară.
Fig. 28 - Sistemul nervos la insecte :
a - văzut ventral; b - văzut dorsal (ganglionii cefalici şi toracici);
Ant - antene; Ao - aorta; Call - corpora allata; Ccar - corpora cardiaca;
Ccen - corp central; Com - comisură; Con - conectiv; Confr - conectiv
frontal; Cped - corp pedunculat; Deut - deutocerebrum; Ghip - ganglioni
hipocerebrali; Gn - ganglioni; Gs - ganglioni subesofagieni; Gven - ganglioni
ventrali; Nant - nervi antenali; Nar - nervii aripilor; Npc - nervii picioarelor;
Nr - nervul recurent; Nsim - nervii sistemului simpatic; Oc - ochi; Ocl - oceli;
Oes - esofag; Pcer - puntea cerebrală; Prot - protocerebrum; Trit - tritocerebrum
(după W e b e r).
35L a n ţ u l n e r v o s v e n t r a l este alcătuit, în general, din 3 perechi de
ganglioni toracici şi 8 perechi de ganglioni abdominali. Acest număr de ganglioni
ai
lanţului nervos ventral se întâlneşte de obicei în stadiul embrionar şi la unele
insecte
inferioare. La marea majoritate a insectelor numărul ganglionilor este mai mic
datorită
contopirii acestora între ei. Astfel, ganglionii toracici fuzionează adesea, uneori
şi cu
prima pereche de ganglioni abdominali, formând o singură masă ganglionară. De
asemenea, ultimele 2 - 3 perechi de ganglioni abdominali se pot contopi între ei.
În
raport cu gradul de fuzionare a ganglionilor la insecte se constată deosebiri
evidente în
privinţa numărului ganglionilor nervoşi, atât de la specie la specie cât şi la
aceeaşi
specie, în funcţie de sex sau stadiul de dezvoltare. Astfel, la larvele de albine
există 3
perechi de ganglioni toracici şi 8 perechi de ganglioni abdominali, pe când la
adulţi
numărul lor se reduce la 2 perechi de ganglioni toracici şi 5 perechi abdominali.
La
unele insecte se pot contopi toţi ganglionii lanţului nervos ventral cu ganglionii
subesofagieni într-o singură masă (la unele homoptere, coleoptere, la larvele unor
diptere etc.). De la ganglionii lanţului nervos ventral pleacă nervi la organele
din
regiunile în care se găsesc. Astfel, ganglionii toracici inervează musculatura
segmentelor respective, picioarele şi aripile, iar ganglionii abdominali
musculatura
abdomenului etc.
S i s t e m u l n e r v o s s i m p a t i c sau v i s c e r a l este alcătuit din
următoarele părţi : sistemul nervos stomatogastric, nervul ventral nepereche şi
sistemul
simpatic caudal.
S i s t e m u l s t o m a t o g a s t r i c este în legătură directă cu
tritocerebrum şi
inervează regiunea anterioară a tubului digestiv (intestinul anterior şi cel
mijlociu).
N e r v u l v e n t r a l n e p e r e c h e, mai dezvoltat în abdomen, se întinde
sub
forma unor fascicule subţiri de-a lungul lanţului ganglionar ventral, printre
conectivele
acestuia. În fiecare segment el prezintă excrescenţe laterale, care inervează
stigmele şi
traheele.
S i s t e m u l s i m p a t i c c a u d a l porneşte de la ultima pereche de
ganglioni abdominali şi inervează regiunea posterioară a tubului digestiv şi
organele de
reproducere.
S i s t e m u l n e r v o s p e r i f e r i c este situat sub hipoderma
tegumentului.
Este alcătuit din numeroase celule nervoase, prevăzute cu nervi care fac legătura
cu
diferite organe şi ţesuturi.
F i z i o l o g i a s i s t e m u l u i n e r v o s. Fiecare ganglion este un
centru
reflex. Prin nervii senzitivi, care leagă organele de simţ de lobul respectiv al
ganglionului, se primesc impulsuri de la periferia corpului, iar de la lobul motor
prin
nervii motori (locomotori) sunt transmise răspunsurile (comenzile) muşchilor
organului
efector. Prin intermediul arcului reflex se realizează legătura între diferitele
părţi ale
corpului şi efectuarea mişcărilor acestuia, sistemul nervos central reprezentând
sistemul
de legătură şi coordonare a mişcărilor insectei.
Creierul, ca principalul centru al activităţii insectelor, coordonează funcţiile
receptorilor optici şi olfactivi. Ganglionii subesofagieni dirijează mişcările
pieselor
bucale. Ganglionii cerebroizi şi cei subesofagieni reglează tonusul muscular.
Sistemul
nervos stomatogastric, prin nervul recurent reglează digestia şi
Willems, P.J.: Genetic causes of hearing loss.

New Engl. J. Med. 342 :1101 – 1109, 2000.
activated protein kinase (2) further activate a
substrate protein. Most protein kinases
phosphorylate tyrosine (tyrosine kinase),
serine, or threonine by transferring a phosphate
NITATE
C
Ă DE BAZĂ
Ă A EVIDE
ENȚEI EC ONOMICE
E
1. Obiective e
1.1

a

C
Contabilitat
s
mult mai
p
dub blă într-o
metrie. Dup
A
Astăzi,
con
c mai comp
o
într-un m
Princip
r reţea de
(după N.
N Feleagă) )
indiferent de
reprezentarea unei
în
economici,
rezultatele
financiare finale.
organizeze şi
Organizarea
dar şi o
organizatoric,
financiar-
contabilă.
cadrul
contabil. În
superioare),
superioare
loc
calendaristic,
calcula diverşi
pozitivi şi
contabilitate
patrimoniului;
lor.

entităţi.

unei entităţi.
13EA DE ÎNV
VĂȚARE 2
UNITATE
OBIEC
CTUL ȘI METODA
M
C
CONTABIL
LITĂȚII
1. Obiective e
2.1
 identi fice obiectu ul de studiu al contabili ității;
 identi fice proced deele ce apar rțin metode i contabilită ății;
 identi fice și să în nțeleagă rolu ul principiil or contabili ității;
 identi fice funcții ile contabil lității și ne ecesitatea ap
plicării ace estora în
contab bilitatea fin anciară și în n contabilita atea de gest tiune.
2 Obiect tul contabil lității
2.2.
C orice di sciplină știi ințifică, con ntabilitatea reprezintă simultan
Ca
s
o teorie
t
și me etodă. În
calitatea a sa de teor rie științific că, contabil itatea repre ezintă un sis stem
de pri incipii și cu unoștințe
care ex xplică și inf formează, iar
i ca meto
odă sau teh hnică, un an nsamblu co oerent de procedee,
p
instrume ente prin ca are se observ vă și înregis strează resu ursele econo omice
ale so ocietății, sep parate ca
utilități patrimonial le.
pții cu
Concep
1. Concep pția admin nistrativă consideră că obiectu ul contabil lității îl
privire la constit tuie reflect area și con ntrolul, în expresie valorică,
v
a faptelor
delimita area admini istrative, în n vederea sp prijinirii ma anagementu ului,
pentru a obține
și defin
nirea cu un minim de eforturi (ch heltuieli) un n maxim de e efecte eco onomice
ului
obiectu (rezult tate-venituri i).
contabi ilității
2. Concep pția juridic că consider ră că obiec ctul contab bilității îl formează
f
patrim
moniul unui i subiect de
d drept (a agent econo
omic), prin n prisma
relațiil or juridice e, adică a drepturil lor și obl ligațiilor p pecuniare
(mater riale) ale unei
u
persoa ane fizice sau juridic ce, în core elație cu

obiecte ele, adică cu
u bunurile și
ș valorile co orespunzăto oare.
3. Concep pția
econ omică
sau u
financia
ară
consid
deră
că
obiectul
contab bilității îl co
onstituie cir rcuitul capit talului, priv vit atât sub aspectul
destina ației lui, câ ât și sub for rma modulu ui de dobân ndire, respe ectiv sub
forma de capital propriu
p
și ca apital străin. .
16 Obiectul contabilității este reprezentat de reflectarea în expresie bănească
a bunurilor mobile și imobile, inclusiv solul, bogățiile naturale,
zăcămintele și alte bunuri cu potențial economic, disponibilitățile bănești,
Concluzie
titlurile de valoare, drepturile și obligațiile agenților economici, precum
și mișcările și modificările intervenite în urma operațiunilor patrimoniale
efectuate, cheltuielile, veniturile și rezultatele obținute de aceștia.
 Obiectivul fundamental al contabilității îl reprezintă furnizarea de
informații utile luării deciziilor menite să asigure o imagine fidelă a
patrimoniului, situației financiare și a rezultatelor obținute.
Din această definiție rezultă că patrimoniul presupune existența a două condiții de
bază: un titular de patrimoniu (subiect de drept) pe de o parte, iar pe de altă
parte bunurile și
valorile economice, privite ca obiecte de drepturi și obligații, precum și
titularul de patrimoniu.
Structura de ansamblu a patrimoniului este următoarea:
PATRIMONIU
BUNURI ECONOMICE
PATRIMONIU PROPRIU
(DREPTURI)
PATRIMONIU STRĂIN
(OBLIGAȚII)
Bunurile economice se concretizează în bunuri materiale (terenuri, clădiri,
utilaje,
mijloace de transport, stocuri de materii prime etc.) şi nemateriale (proiecte,
programe
informatice etc.) formând averea entității.
Relaţiile de drepturi se referă la faptul că entitatea îşi procură o parte din
avere din
surse proprii, iar bunurile procurate îi aparţin de drept.
Relaţiile de obligaţii se referă la faptul că entitatea îşi procură o parte din
avere din
surse împrumutate, fiind obligată să restituie terţilor contravaloarea bunurilor
procurate.
Contabilitatea se ocupă cu reflectarea în expresie valorică a patrimoniului, ea
înregistrează circuitul elementelor patrimoniale în condiţii concrete de timp şi
spaţiu,
calculează mărimea acestor elemente şi reflectă mişcarea patrimoniului prin
operaţiuni de
intrări şi ieşiri 1 .
1
Ghe. Talaghir, Ghe. Negoescu – Contabilitatea pe înţelesul tuturor. Editura All,
Bucureşti, 1998.
17Contabilitatea studiază modul de gestionare a patrimoniului, fundamentează
deciziile referitoare la finanţarea şi utilizarea elementelor patrimoniale,
controlează realizarea
deciziilor şi stabileşte răspunderi privind integritatea şi dezvoltarea
patrimoniului.
De asemenea, contabilitatea studiază echilibrul global al patrimoniului, prin
respectarea ecuaţiei patrimoniale:
BUNURI ECONOMICE = DREPTURI + OBLIGAȚII
cu derivatele sale:
Drepturi = Bunuri economice – Obligaţii
şi
Obligaţii = Bunuri economice – Drepturi
În consecinţă, entitatea reprezintă o unitate patrimonială, al cărei patrimoniu
poate fi
privit sub dublu aspect: al mijloacelor economice (bunurile/averea) şi al surselor
de
procurare a acestor mijloace (capitalul) proprii şi străine.
Mijloacele economice definesc activul patrimonial, iar sursele definesc pasivul
patrimonial.
În derularea operaţiunilor economico-financiare, apar şi o serie de procese
economice, sub forma veniturilor şi cheltuielilor, care ajută la înregistrarea
creşterii sau
diminuării patrimoniului.
În acest context, ecuaţia patrimonială de mai sus, devine:
AVERE = CAPITAL
182.3. Metoda contabilității
Datorită complexității obiectului de studiu, metoda contabilității reunește mai
multe
procedee tehnice de lucru.
Un ansamblu de procedee aflate într-o strânsă corelație și
Metoda intercondiționare ca un tot unitar, în vederea stabilirii normelor și
contabilității principiilor cu caracter special pe care se fundamentează
contabilitatea
și cu ajutorul cărora cercetează starea și mișcarea elementelor
patrimoniale ale unităților patrimoniale.
O trasătură caracteristică a metodei contabilității este aceea a folosirii unor
procedee
care să permită înregistrarea numerică, cifrică, a existenței și mișcării
patrimoniului unităților
economice și sociale în expresie valorică. Generalizând, se poate spune că metoda
contabilității reprezintă totalitatea procedeelor interdependente, pe care le
folosește aceasta în
scopul cunoașterii situației patrimoniului și a rezultatelor obținute.
1. procedee comune tuturor științelor;
Procedee utilizate 2. procedee specifice metodei contabilității;
de contabilitate 3. procedee ale metodei contabilității, comune și altor discipline
economice.
Procedee comune Observația – este faza inițială a cercetării obiectului de studiu
al
tuturor științelor oricărei științe și este utilizată pentru cunoașterea
operațiilor economice
care se pot exprima valoric și pe care le reflectă cifric, numeric, cu
ajutorul procedeelor sale specifice.
Raționamentul – se aplică de metoda contabilității, pentru că pe bază
de judecăți logice, pornind de la fenomenele și procesele economice
care intră în obiectul său de studiu, să ajungă la concluzii noi (ex.:
activul este egal cu pasivul, pentru că între mijloacele economice și
sursele de finanțare a acestora există o egalitate perfectă).
Comparația – se folosește de metoda contabilității prin alăturarea a
două sau mai multe fenomene și procese economice care se pot exprima
19valoric, cu scopul de a stabili asemănările și deosebirile dintre ele, ca
astfel să se tragă o serie de concluzii. Se folosește frecvent pentru a se
compara veniturile și cheltuielile pe baza cărora se stabilesc rezultatele
finale.
Clasificarea – acțiunea de împărțire, distribuire, repartizare sistematică
pe clase sau într-o anumită ordine a obiectelor în funcție de asemănările
și deosebirile dintre ele. Asemănările le apropie și le încadrează în
aceeași clasă, iar deosebirile le diferențiază și le distribuie în clase
diferite.
Analiza – procedeu științific de cercetare a unui întreg, a unui fenomen
care se bazează pe examinarea fiecărui element component în parte.
Sinteza – ca procedeu științific de cercetare a fenomenelor se bazează
pe trecerea de la particular la general, de la simplu la compus pentru a
se ajunge la generalizare.
Procedee specifice Bilanțul – stă la baza dublei reprezentări a patrimoniului în
metodei contabilitate. El furnizează informații generale privitoare la situația
contabilității economică și financiară a unei entități, dar reflectă și la
relațiile ei
economice cu alte entități, fiind completat de o serie de situații anexă
prin care se explică și se detaliază anumite laturi ale activității
economico-financiare ale societății.
Contul – se deschide în contabilitatea curentă pentru reflectarea
fiecărui element din patrimoniu. Contabilitatea dispune de un sistem
de conturi în care reflectarea operațiilor economice rezultate din
mișcarea elementelor patrimoniale are la bază dubla înregistrare.
Balanța de verificare – asigură în contabilitatea dublei reprezentări și
a dublei înregistrări, garanția exactității înregistrărilor efectuate în
conturi. Datele balanței de verificare stau la baza întocmirii bilanțului.
Balanța de verificare îndeplinește atât o funcție de control, cât și o
funcție economică, constituind puntea de legătură între cont și bilanț.
20Procedeele metodei Documentația – orice operație economică și financiară
referitoare la
contabilității comune existența și mișcarea elementelor patrimoniale trebuie să fie
și altor discipline consemnată în documente care fac dovada înfăptuirii lor.
economice Evaluarea – procedeul prin care datele contabilității sunt reprezentate
printr-o singură unitate de măsură, creând posibilitatea centralizării
lor cu ajutorul balanțelor de verificare și generalizarea cu ajutorul
bilanțului. Evaluarea constă în transformarea unităților naturale în
unități monetare cu ajutorul prețurilor.
Calculația – este strâns legată de evaluare ca procedeu al metodei
contabilității. Acest procedeu își găsește aplicarea cea mai largă în
domeniul calculației costurilor de producție.
Inventarierea – se folosește pentru a se cunoaște situația reală a
patrimoniului reflectat în contabilitate, și trebuie să se verifice
existența faptică a tuturor elementelor sale, în scopul descoperirii
neconcordanțelor dintre datele înregistrate în conturi și realitatea de
pe teren.
2.4. Principiile contabilității
Pentru a se oferi o imagine fidelă a patrimoniului, a situației financiare și a
rezultatelor
obținute de către entitate, trebuie respectate cu bună credință regulile privind
evaluarea
patrimoniului și celelalte norme și principii contabile.
Principiul  nu sunt admise supraevaluarea elementelor de activ și a
prudenței veniturilor, respectiv subevaluarea elementelor de pasiv și a
cheltuielilor, ținând cont de deprecierile, riscurile și pierderile
posibile generate de desfășurarea activității unității;
 prudența presupune anticiparea efectelor unor acțiuni și în special
a transferului de proprietate cu efecte posibile asupra exercițiului
sau a celor parcurse deja, întrucât asupra lor nu se mai poate
interveni din punct de vedere contabil;
 la încheierea fiecărui exercițiu financiar se contabilizează
datoriile și pierderile probabile.
21Principiul
 continuitatea aplicării regulilor și normelor privind evaluarea,
permanenței înregistrarea
în
contabilitate
și
prezentarea
elementelor
metodelor patrimoniale și a rezultatelor asigurând comparabilitatea în timp a
informațiilor contabile;
 asigură aplicarea pentru aceleași elemente, stucturi, domenii de
activitate, a acelorași metode de la un exercițiu la altul, excluzând
schimbarea metodelor în cursul exercițiului;
 modificarea metodelor de la un an la altul trebuie să fie
determinată de o profundă motivație (modificarea unor acte
normative, stabilirea unor reguli generale de evaluare noi etc.).
Principiul
 presupune că unitatea patrimonială își continuă în mod normal
continuității funcționarea într-un viitor previzibil, fără a fi în stare de
lichidare
activității sau de reducere sensibilă a activității;
 atunci când funcționarea este delimitată în timp sunt menționate
datele de începere și de încetare a activității;
 permite fixarea responsabilității entității
residue from adenosine triphosphate (ATP),
which is then converted to adenosine di-
phosphate (ADP). Other receptors mediate the
removal of phosphate from a phosphorylated
tyrosine side chain by means of their
phosphatase activity (3). With one important
type of receptor, ligand binding activates
guanylate cyclase (4), which catalyzes the for-
mation of cyclic guanosine monophosphate

(cGMP) from guanosine triphosphate (GTP).
The cGMP functions as a second messenger and
brings about a rapid change of activity of
enzymes or nonenzymatic proteins. Removal or
degradation of the ligand reduces the conc
In Figure l(b), u(xo,t) is the vertical displacement, at time t, of the point on

the string
that would be at Xo on the x-axis, if the string were in the rest position of
Figure l(a). By
assumption of transverse vibrations, there is no horizontal displacement. The
vector TR(xo,t)
(cf. Figure 2( a)) is the force that the portion of string to the right of Xo
exerts on the portion to
the left of xo' We call TR the right-tension at xo' at time t. Similarly, we define
the
left-tension TL(xo,t). We assume that TR(xo,t) is tangent at Xo to the graph of
u(x,t), as a
function of x, and we assume that TL(xo,t) = -TR(xo,t). This last assumption might
seem to
imply the points of the string will never move, since the net force on each point
is TL + TR = O.
However, since the mass of a point is 0, the acceleration of a point is not
determined by
Newton's equation F = rna. To obtain the acceleration of a point x o, we need to
take the limit
of the ratio of the net force on the portion of the string above the interval [xo-
h, Xo+h] to the
mass of this portion, as h .... 0 (cf. Figure 2( a)).
u
u
(a)
(b)
Figure 2
We compute this limit as follows. Let the magnitude of the left(or right)-tension
at Xo at time
t, be denoted by T(Xo,t). This is called the tension at Xo at time t. We assume
that the
tension is the positive constant To, when the string is straight (as in Figure
l(a)). If the string is
stretched by a factor of s , relative to its length at rest, then the new tension
will be of the form284
Chapter 5
The Wave Equation
g(s)'To ' for some function g(s), which depends on the nature of the string. That
is, we are
assuming that the function g depends only on s, and not on other quantities such as
the rate of
stretching or temperature variations due to bending. Of course, g(l) = 1 , because
when the
string has not been stretched (Le., s = 1), the tension is To. We assume that g(s)
is C1 for
stretches s which are encountered during vibrations. Using these assumptions
together with the
restriction that the vibrations are transverse, we will show that Newton's second
law impies that
g(s) = s (Le., the tension in the string is proportional to the stretching factor
s). Then we will
see that the PDE for u(x,t) which results (without any of the usual dubious
statements about u~
being negligible) is necessarily a linear PDE, namely the wave equation.
Definition. A string is linearly elastic for a ~ s ~ b, provided that, when the
string, at tension
To, is stretched by a factor of s in [a,b], the tension becomes Tos (Le., g(s) =
s).
More precisely, we will show that, in conjunction with Newton's law, the assumption
that the
string admits nontrivial transverse vibrations actually implies that the string is
linearly elastic for
s in any range which is encountered during such a vibration. In Figure 2(b) above,
the string in
the interval [x,x + box] has been stretched by about a factor of (box 2 + bou 2)!
jbox which tends
to s == (1 + u~)!, as box -+ o. The direction of the right tension at x is
approximately the same
as that of the unit vector (boxi + bouj) j (box 2 + bou 2)!, which approaches the
exact unit tangent
vector (i + uJ)j(1 +
u~)! = ~ (i + uxj) ,as box -+ o. Thus,
Consequently, the net force on the string between x - h and x + h, as in Figure
2(a), is
Since we are considering only transverse vibrations, the mass of the portion of the
string, from
x-h to x+h, remains the same as it would be if the portion were in the straight
rest position,
namely 2hD, where D is the mass per unit len~th of the string when straight (Le., D
== the
linear density of the straight string in Figure l(a»). By Newton's law and the
assumption of
purely transverse vibrations, the acceleration of the string at x is
= u 1 ox.
a
Equating the
[T R ]
= U 1 ox.
a
[T 0 g{§l].
+ U
1
S
1
and j components of both sides, we have
a [T
ox.
0
g{§l
s U x ]. JSection 5.1
0= (To/D)
and
Utt =
where s
285
Thf Wave Equation - Derivation and Uniqueness
~ [¥)]
(I)
(To/D).~uxx + (To/D)'~ [~]
(J)
ux '
= (1 + u;)t . Thus, (I) implies that G(s) == g(s)/s is independent of x. Now
(I' )
for all Xl and x2 in [O,L] (Why?). For any fixed time t, we may choose xl to be a
point such
that
UX(xl,t)
is
O. (Here we assume that the ends of the string are fixed and apply Rolle's
theorem.) Thus S(XI,t) = 1, and since G(I) = g(I)/1 = 1, (I') yields 1 = G(S(x2,t))
for any
x2 and any t during the vibration. In other words, g(s) = s for any stretch s which
is
encountered during a transverse vibration. Thus, the PDE (J) reduces to the wave
equation
In many derivations, the PDE (J) is linearized (cf. Section 1.2) by the outright
that u x 2 is negligible, so that one can (all too conveniently!) set s = 1 in the
PDE
assumption
.
Remark.
(J), which then becomes the linear wave equation (**). However, the assumption that
u; is
small presupposes some knowledge about the solution of the typically nonlinear PDE
(J) (cf.
Example 11 of Section 1.2, for the pitfalls of linearization). Rather than
introducing this
questionable assumption, we have shown that it is unnecessary to do so, by
demonstrating that
g(s) = s follows from the assumption of transverse vibrations and Newton's law. The
property
g(s) = s means that the string behaves as a spring or a rubber band which is
stretched, but not
by too much, so that Hooke's law holds (cf. Example 6 of Section 1.1). However, not
all strings
satisfy g( s) = s, except in a very short interval about s = 1. For example, the
tension in a piece
of twine can increase enormously even for small stretches, whereas for a string of
taffy the tension
may even eventually decrease upon stretching. In essence, we have shown that such
strings do not
admit purely transverse vibrations u(x,t) for which the stretching factor, (1
outside the range of linear elasticity for some (x,t).
+ ux (x,t)2) t,
lies
0
Solving the standard initial value problem for a string with fixed ends
As with the heat equation, we first find all of the product solutions of Utt
method of separation of variables. Substituting u(x,t)
X(x)T"(t)
= a 2X"(x)T(t)
or
= X(x)T(t)
into Utt
= a 2uxx ' using the
= a 2uxx ' we get
~ = ~ = c = ±,A2
a 2 T(t)
X(x)
,Chapter 5
286
The Wave Equation
where A is some nonnegative constant. The ODE X" = ± A2X for X(x) is exactly the
same as
for the heat equation in Section 3.1, but the solutions of the ODE Til = ±A 2a2T
for T(t) are
different, because of the second time derivative. The possible product solutions
fall into the
following three cases, where cl ' c 2' d l and d 2 are arbitrary constants:
Case 1 (c
= _A2 < 0):
(1)
Case 2 (c = A2 > 0):
u(x,t) = (dle Aat
+ d 2 e -Aat H cle AX + c 2 e -AX).
(2)
Case3 (c = A2 = 0):
(3)
We will now formulate one of the simplest standard initial/.boundary-value problems
for
the wave equation. Recall that in solving Newton's equation mx '(t) = Flt), it is
necessary to
specify x( to) and x I (to) in order to obtain a unique solution for the position
x( t) of a particle.
For the wave equation (whose derivation was based on Newton's equation), it is also
necessary to
specify not only the initial profile u(x,O) of the string, but also the initial
velocity ut(x,O).
Otherwise, we will not obtain a unique solution u(x,t). First we assume that the
ends of the
string are fixed (Le., u(O,t) = 0, ulL,t) = 0), although in Section 5.3, we
consider the cases
where one or both of the ends are allowed to slide vertically. We expect that under
reasonable
circumstances the following standard problem will have a unique solution:
°
2
= a uxx '
~ x ~ L,
u(O,t) = 0, u(L,t) = 0,
D.E.
B.C. U tt
I.C. {
u(x,O)
= f(x),
ut(x,O) = g(x),
--00
< t <
+00 ,
(4)
(initial position)
(initial velocity).
We require that u(x,t) have a C2 extension to an open domain that contains the
strip
~ x ~ L, --00 < t < 00. As with the heat equation, the only product solutions which
satisfy the
and A = n7r/L, n = 1, 2, 3, .... By taking linear
B.C., are of the form (1), where c 2 =
combinations of such solutions of the D.E. and B.C., we obtain a solution of the
form
°
°Section 5.1
287
The Wave Equation - Derivation and Uniqueness
(5)
Note that
(6)
Substituting t =
° in (5) and (6) yields
(7)
In summary, we have shown the following result.
Proposition 1. A solution of the problem
°
D.E.
B.C.
~ x ~
u(O,t)
= 0,
u(L,t)
L,
-w
< t < +00 ,
= 0,
(8)
N
u(x,O) = f(x) = lnN=lBnSin(n11X/L),
I.c.
1
ut(x,O) = g(x) = t=l A nsin(n11X/L)
is
(9)
Remark. Here we have expressed u(x,t) in terms of the Fourier sine coefficients Bn
and An of
f(x) and g(x) respectively. Note that An in (5) is (L/n7ra)An .
0
Of course, one can pose problems where f(x) and g(x) are not finite sine series as
above.
But if f(x) and g(x) are continuous and piecewise
on rO,L), with f(O) = f(L) =
and
g(O) = g(L) = 0, then we know that f(x) and g(x) may be unitormly approximated to
within any
small positive error by partial sums of their Fourier sine series (d. Theorem 1,
Section 4.3). Thus,
such f(x) and g(x) can be replaced by finite sine series within experimental error.
Later we will
show that two solutions must be close, if the initial profiles and velocity
distributions of the two
solutions are close (d. the Corollary to Theorem 5 in Section 5.2).
c1
°Chapter 5
288
The Wave Equation
Example 1. Solve the initial/boundary-value problem
°
2
= a U xx '
~ x ~ L,
u(O,t) = 0, u(L,t) = 0,
D.E.
B.C. U tt
I.C. [
u(x,O) = f(x) =
-00
< t < +00,
2sin(~),
ut(x,O) = g(x) = sin(¥) -
3sin(~)
Solution. We simply apply Proposition 1 with B3 = 2,
.
Al = 1, A5 = -3, and with all of the
other An and Bn equal to zero. Then
Note that to get the solution quickly, one can simply insert a factor of
cos{mrat/L) in the terms
involving sin t n1lX{L) in f(x) and insert a factor of (L/mra)sin(n7rat/L) in the
terms involving
sin(n1lX/L) in g(x, and then add the results to get u(x,t). 0
Harmonics
The individual terms of the series (5), namely (for n = 1,2,3, ... )
un ( x,t ) =
· (n7rat)
[A nsm
-r- + Bncos (n7rat)]
-r- sm . (n1lX)
L
are called the harmonics or overtones of the string with fixed ends at x =
These constitute a complete family of product solutions of the D.E.
U tt
°
(10)
and x = 1.
= a 2u xx with B.C.
u(O,t) = u{L,t) = 0, as n runs through the values 1,2, .... If An and Bn are not
both zero,
we can rewrite un(x,t) as follows. Let Rn:: (A; + B;)'t.
= An/Rn and sin{ On) = Bn/Rn ,since (An/Rn)2 + (Bn/Rn)2 =
un(x,t) =
Then there is a On' such that cos{On)
1. Thus,
Rn[cos(On)sin{~) + sin(On)cos(~)]sin(T) = Rnsin(~ + 0n)sin(T) .
We see that un(x,t) oscillates between ±Rnsin{n1lX/L) as t varies. Rn is called the
amplitude
of un(x,t) and On is the phase of un(x,t) (cf. Figure 3, where Rn= 1, L =
6 ).Section 5.1
u
u
u
289
Figure 3
The time that it takes a harmonic to complete one oscillation is called its period.
The period is
inversely proportional to n. To find the period of the n-th harmonic, we set
ll1rat/L = 211" and
solve for t, obtaining t = 2L/na. The frequency of a harmonic is the number of
oscillations per
unit time, and it is just the reciprocal of the period. Hence, the period and
frequency of the n-th
harmonic of the string (with fixed ends) are respectively given by
p
n
= na
2L
and
1
na
vn = p = 2L.
n
(11)
Uniqueneljs and the energy integral
As with the heat equation, there is the uniqueness question.
solutions to problem (4) for a given f(x) and g(x)?
Can there be two different
Theorem 1 (Uniqueness). Let u 1 (x,t) and u 2 (x,t) be C 2 solutions ofthe
following problem
D.E. U tt = a 2 uxx '
$ x $ L, -00 < t < +00,
°
= A(t) , u(L,t) = B(t) ,
u(x,O) = f(x) , ut(x,O) = g(x) .
B.C. u(O,t)
I.C.
Then u 1 (x,t) = u 2 (x,t) for all
° $ x $ L,
-00
< t < +00.
Proof. Let v(x,t) = u 1 (x,t) - u 2 (x,t). Note that v satisfies the related
problem with
and vt(x,O) = 0. We need to show that
homogeneous B.C. and I.C .. In particular, v(x,O) =
v(x,t) = 0 for all t. In the proof of the corresponding Theorem 1, in Section 3.2,
we
°
accomplished this by proving that
Here, we will prove that the function
J:
[v(x,t)] 2 dx =
° via differentiation with respect to
t.Chapter 5
290
The Wave Equation
(12)
is zero. Once this is done, then vt(x,t) =.0 for all x in [O,L] and all real t, and
(as required)
L
t
v(x,t) = v(x,t) - v(x,O) =
vt(x,t) dt = 0.
We compute H'(t), using Vtt = a 2vxx and differentiating under the integral (cf.
Appendix A.3) :
H'(t) =
J:
= 2a
[a22vxvxt + 2v t v tt ] dx = 2a2
2f L 0 ox
o (vxvt ) dx
J:
[VXVxt + vtvxxl dx
= 2a 2 [vx(x,t)vt(x,t)]
IL
0
= 0,
°
since (by the B.C. v(O,t) = and v(L,t) = 0) vt(O,t) =~ v(O,t) = 0, and similarly
vt(L,t) = 0.
Since H' (t) = 0, we know that H(t) is constant, but this constant is 0, since H(O)
=
according to the initial conditions for v (note that vx(x,O) = ~ v(x,O) = 0). 0
°
Remark. The function H(t) (cf. (12)) in the above proof is actually proportional to
the total
energy of the string given by v(x,t). Indeed, the kinetic energy of the segment of
the string from
x to x + ill is approximately !(DLlx)[ut (x,t)]2, where D is the mass per unit
length. The
work done (energy expended or potential energy) in stretching the segment from Llx
to
(Llx2+Llu2)t ~ [1 + ux (x,t)2]tLlx = sLlx , is the following integral of the force
(tension) with
respect to the increase r in the length of the segment during the stretching
process.
f °
(S-I)LlX
I
Llx+r
r 2
(s-I)Llx
To ----x-::- dr = To·(r + O"X"":":")
uX
~uX
°
= ToLlx(s -1 + !(s _1)2) = !To(s2 -1)Llx
= ! T o[ux(x,t)12 ill ,
where we have assumed, as in the derivation of the wave equation, that the string
is linearly
elastic, with rest tension To. Hence, the potential energy of the segment of string
from x to
x + Llx is ~ !To[ux (x,t)]2 Llx. When this is added to the kinetic energy and the
result is
integrated, we get the energy integral (or simply, energy) of the string at time t
E(t)
(13)Section 5.1
291
°
which is tD times H(t) in (12), using v for u. For an unforced string, we expect
E(t) to be
constant. Thus, it is not surprising that H/(t) =
in the preceding proof. Of course, the
physical intuition is no substitute for the explicit computation of H' (t) above,
but it does lead us
to consider the function H( t) in the first place. In other words, the physical
basis for the proof of
the uniqueness (Theorem 1) is the law of conservation of energy. 0
Example 2. Calculate the energy of the n-th harmonic
' (n1l'at)
u ( x,t ) = [A nsm
----r;-
. (n1l'X)
+ Bncos (n1l'at))
----r;- sm
T
.
Solution. The energy E(t) is defined by (13), and we know from the proof of Theorem
1 that
E(t) is constant. Thus, E(t) = E(O), and so we only need to compute E(O):
E(O) =
~ J~
=:2 1
[T o [U x (x,0))2
+ D[ut (x,0)]2]
dx
JL 0 [ TO[L
n 11' Bncos(T)]
n
2 + D[T
n 1I'a Ansm(T)]
. n
2] dx
1I'X
1I'X
L
=
L
°
°
~(!!.[)2 [ToB~ J cos 2 (T) dx + Da2 A~ J sin2 (T) dX]
n 2 11'2
2
2 2
2
11'2
2
2
11'2
2 2
="""4L (ToBn + Da An) = 4L To (Bn + An) n = 4L ToRn n ,
where we recall that a
==
.I.
2
2
(To/D)2 and Rn = An
.
+ Bn 2. IS the square of the amplItude
of the
harmonic. It is possible, but rather lengthy, to compute E(t) directly by
considering [ux (x,t))2
and [ut(x,t)f, Of course, we know that the end result will be E(O) anyway.
0
Remark. Note that the energy of the n-th harmonic is proportional to n2 for a given
amplitude,
and it is proportional to the tension and the square of the amplitude. However,
unlike the
frequency /In = tnLvT;JIT, the energy is independent of the linear density D. 0
Example 3. (The motion of the plucked string) Find the formal solution of the
problem for the
motion of the plucked string:
D.E.
U tt
2
= a uxx '
o ~ x ~ L,
B.C. u(O,t) = 0, u(L,t) = 0,
I.C.
u(x,O)
= f(x),
ut(x,O)
= 0,
where f(x) is the function with graph shown in Figure 4.
-00
< t < 00,
(14)Chapter 5
292
The Wave Equation
U
U
o ------------------------------------------------:
:
!
!
!
! !
o
x
L
(uo > 0, 0 < Xo < L)
Figure 4
(Le., the string is "plucked" at some fixed Xo in (O,L), lifted to the displacement
released with zero initial velocity, Le., ut(x,O)
= 0).
Uo and
Solution. We have
o ~ x ~ x o'
Xo
~
x
~
L.
The formal solution of problem (14) is obtained by computing the Fourier sine
coefficients Bn of
f(x) (cf. Proposition 1) and using formula (9) with N
=
00.
(The notion of a formal solution was
introduced in Section 4.3.) Note that f is continuous and piecewise C 1.
Integration by parts is
valid for such functions. Thus,
The endpoint evaluation is zero, since f(O) = f(L) = 0 , and from the fact that
o ~
Xo
we obtain
Bn =
=
f ;11"[
to ~
o
0
cos(nr) dx
[u . (nn)
r 2 [ n L]2
11"
~ sm -,:;-0 -
2
+
u
The formal solution is
Xo 0
T
< x
~
L,
cos(nr) dX]
. (nn)]
~r sm
_
2L uo_ 1 sl·n(n nO)
-2
,-,:;-.
11" Xo (L-xo) n
r ~r
< Xo
x
2Lu o [1
= ~ Xo
1 ] . (nn
+ L-Xo
sm -,:;-0)Section 5.1
293
(15)
Let uN(x,t) denote the sum of the first N terms of(15).
Then uN(x,t) is a COO solution of the
D.E. with the given B.C .. Moreover, uN(x,O) is the N-th partial sum SN(x) of FSS
f(x), and
thus we know that for N sufficiently large uN(x,O) will approximate f(x) to within
any
preassigned error (cf. Theorem 1 of Section 4.3). Hence, for practical purposes,
the problem of the
plucked string is solved. While it is possible to prove that the sum (15) converges
(i.e., u(x,t) is
defined for all (x,t)), u(x,t) is not C2 and hence is not a strict solution of the
D.E .. Using the
techniques of Section 5.3, one can show that for most times t, the graph of u(x,t)
in (15) consists
of three line segments whose slopes do not match at the two interior corners where
pairs of them
join. Indeed, the two corners move in opposite directions (and with horizontal
speed a) around
the parallelogram formed by the original profile and its reflection through the
point tL on the
x-axis (cf. Figure 5 below). Observe that u(x,t + 2L/a) = u(x,t). In particular,
the string
returns to the ori,ginal plucked position at t = 2L/a and repeats its motion. The
values L/n,
2L/n, 3Ljn, ... , (n-l)L/n, where sin(n11X/L) vanishes are called the nodes of the
harmonic
cos(n1rat/L)sin(n11X/L). Formula (15) shows that the harmonics, with Xo as a node,
drop out of
the sum, because of the factor sin(n11Xo/L).
0
u
.......... ~ ....... ..
.
', . . . ".! ~
j.-at --k-at -..i
0 ", .
........
"
.........
.... " ......
.........
Figure 5
xChapter 5
294
The Wave Equation
Summary 5.1
1. Derivation of the wave equation: Under the assumption that a string of linear
density D,
stretched with tension To between two points, is executing only transverse
vibrations, we
demonstrate (via Newton's equation F = rna) that the amplitude u(x,t) of the
vibration must
obey the wave equation U tt = a2uxx ,where a2 :: TolD. In the course of the
derivation, we
prove that a string which undergoes transverse vibrations must be linearly elastic
(Le., the tension
at any point is of the form sT o ' where s:: (1 + u~)t is the local stretching
factor).
Consequently, it is not necessary to assume that u~ is negligible in order to
achieve a linear wave
equation for u(x,t), for transversely vibrating strings.
2. The standard problem for fixed ends (Proposition 1): A solution of the problem
°
D.E.
B.C.
~ x ~ L,
u(O,t)
= 0,
u(L,t)
-00
< t < +00 ,
= 0,
(81)
N
u(x,O) = f(x) = lnN=1 Bnsin(T)'
I.C.
[
ut(x,O) = g(x) =
~n=1 Ansin(T)
(82)
is
3. Harmonics: The product solutions of the D.E. and B.C. of (81) are called
harmonics and they
are of the form (where n = 1,2,3, ... )
where Rn = (A;
+ B;)t is the amplitude of the harmonic, and
define the phase On'
An/Rn = cos On' Bn/Rn = sin OnSection 5.1
295
4. Energy: The energy at time t of any solution u(x,t) of (SI) is given by
IJL [ To[ux(x,t)] 2 + D[ut(x,t)] 2] dx,
E(t) ="2
(S4)
0
which was shown to be constant in the proof of the uniqueness theorem (Theorem 1).
The energy
of the harmonic un(x,t) in (S3) was computed to be
2
it To R~ n2 in Example 2.
5. Uniqueness: Theorem 1 implies that (S2) is the unique solution of the problem
(SI). More
generally, even if the B.C. in (SI) are replaced by u(O,t) = A(t) and u(x,t} = B(t)
and if f(x)
and g(x} are not necessarily finite Fourier sine series, Theorem 1 states that
there is at most one
(possibly no solutions) C2 solution of the resulting problem. The proof proceeds by
showing that
the energy of the difference of two solutions is time-independent and initially
zero.
6. The plucked string:
The formal solution for the problem of the plucked string is found in
Example 3. By truncating the formal solution, one can produce COO solutions of the
D.E. and B.C.
which meet the I.C. to within any preassigned experimental error. However, it can
be shown that
the full sum of the formal solution converges to a function (which is not even C 1)
with corners
which move in opposite directions around a parallelogram, as in Figure 5.
Exercises 5.1
1. Solve the problem
D.E.
U tt
2
= a uxx '
o S x S L,
~
< t < 00,
B.C. u(O,t) = 0, u(L,t) = 0,
I.C. u(x,O)
= f(x),
ut(x,O)
= g(x).
in the following cases:
(a) f(x) = 3sin(![) -
(c) f(x) = 0,
g(x)
sin(!p), g(x)
= ~sin(~),
= sin(![)cos 2 (![),
(d) f(x)
(b) f(x)
= [sin(![)]3,
= [sin(![)]3,
g(x)
g(x)
= 0,
= sin(![)cos2(![).
Hint. For (b) and (c), use trigonometric identities. For (d), use the superposition
principle.
2. Suppose u(x,t) solves U tt = a2uxx ' (a f 0).
(a) Let a, (3, Xo and to be constants, with a f
v(x,t)
= u( ax+x o,(3t+t o), satisfies
v tt
o.
= ((3a/0')2 v xx .
Show that the function v(x,t), given byChapter 5
296
The Wave Equation
(b) For any constant w, let x = cosh(w)x + a·sinh(w)t and t = a-1 .sinh(w)x +
cosh(w)t.
Recalling that cosh 2(w) - sinh2(w) = 1, show that x = cosh(w)x - a,sinh(w)t
t = _a-I. sinh( w)x + cosh( w)t (Le., (x,t) -+ (x,t) is an invertible change of
variables.)
(c) Define u(x,t)
= u(x,t).
Show that Utt - a 2u xx
Ux = uxxx + uttx =
Remark. The transformations (x,t)
uxcosh(w)
= U tt -
+ u t a-1sinh(w), and compute
-+
and
a 2u xx ' mnt. By the chain rule,
Uxx and Utt similarly.
(x,t), known as Lorentz transformations, mix space and
time. Part (c) shows that the wave equation Utt - a 2uxx = 0 retains its form under
Lorentz
transformations. In this way, Albert Einstein (1879-1955) was led to his famous
unification of
space and time (Le., relativity). In most physics books, cosh(w) is written as
(1_v2/a2 )-t, and
sinh(w) is then
and a
:I:
i'(1-v 2/a 2 )-t (Why?), where vi is the relative velocity of two observers,
= C:: the speed of light
Rj
2.99
x
108 m.s-1 .
3. In the derivation of the wave equation (Section 5.1) we did not consider the
effect of gravity
which exerts an additional force of -2hDg j
Xo
+ h, where
on the segment of string between
Xo - hand
g = 32 ft/sec 2 is the acceleration due to gravity.
(a) Deduce that u(x,t) obeys Utt
= a2uxx -
g , if the effect of gravity is considered.
(b) Find a solution of Utt = a 2u xx - g which satisfies the B.C. u(O,t) = 0 and
u(L,t) = 0 and
which is time-independent [Le., u(x,t) = U(x)]. What does this solution represent?
4. In Section 5.3, we show that the B.C. ux(O,t) = 0 means that the end x = 0 of
the string is
free to slide vertically (and similarly, for the end x = L). Show that the proof of
Theorem 1
yields uniqueness in the cases when one or both ends are free to slide.
5. Let v(x,t) and w(x,t) be two C2 solutions of the problem
D.E. Utt
= a 2 uxx'
0 ~ x ~ L,
-00
< t < 00,
B.C. u(O,t) = 0, u(L,t) = O.
(a) Use the technique in the proof of Theorem 1 to show that
~
mnt. vxtwx
f:
[a2vx (x,t)wx (x,t)
+ vt(x,t)wt(x,t)]
+ vxw xt + vxxw t + vtwxx = (vtwx + vxwt)x.
dx
= O.Section 5.1
(b) Let B(v,w) =
J:
297
[!Tovxwx + !Dvtw t ] dx.
In part (a), we proved that B(v,w) is a constant, independent of t. Note that the
energy Eu of
u(x,t) is B(u,u). Establish the result
B(v+w,v+w) = B(v,v) + B(w,w) + 2B(v,w).
(Le., Ev+w :f: Ev + E w' unless B(v,w) = 0).
Hint. Rather than writing out B(v+w,v+w) in terms of an integral, note that B
clearly has the
properties B(u t ,u 2) = B(U2'U t) and B(u t +u 2,u) = B(ut,u) + B(u 2 ,u), which
are all that is
needed to get (*).
(c) Show that for any two harmonics, say un(x,t) (formula (10)) and um(x,t) where m
f=
n, we
have B(um,u n) = 0. Conclude from (b) that the energy of um + un is the sum of the
energies of
un and u m . Is this still true if n = m? Why not ?
(d) Show that the energy of
2:Jn~a Ansin(~)
u(x,t) =
t
2
N
Eu = 7r 4 o 2n=t
N
Hint. Eu
= B(u,u) = B(2n=tUn
N
n2[B~ +
+
BnCOS(~)]
is
(LA n/n7ra)2].
N
, 2n=tUn)
sin(T)
N
= 2n=t B(un,un) +
2 2B(um,un)
= t=tB(un,un).
m<n~N
6. Consider the problem
D.E.
°
= U xx '
B.C. u(O,t) = 0,
I.C. u(x,O) = x( 7r -
U tt
~ x ~
u(7r,t)
7r,
-00
< t < 00,
= 0,
x), ut(x,O)
The Vaccinium genus contains several valuable fruit and ornamental species, among
others: highbush blueberry (Vaccinium×corymbosum L.), cranberry (Vaccinium
macrocarpon Ait.), and lingonberry (Vaccinium vitis-idaea L.). In some most popular
and valuable cultivars, the conventional propagation methods, exploiting hard or
soft wood cuttings, are inefficient. The demand for nursery plants could be
fulfilled only by micropropagation. In principle cultivars are propagated in vitro
through similar three- stage method, based on subculture of shoot explants on
different culture media supplemented with IAA (0–4 mg/L) and 2iP (5–10 mg/L), and
rooting shoots in vivo. The obtained plantlets are transferred to peat substrate
and grown in the glasshouse until the end of growing period. The development of
adventitious shoots should be monitored and controlled during in vitro stages.
Many clones have specific require- ments for growing conditions and/or are
recalcitrant.
Key words: Adventitious shoots, Axillary shoots, Blueberry, Cranberry, Lingonberry,
Nodal explants
1. Introduction
Highbush blueberry (Vaccinium×corymbosum L., syn. Vaccinium corymbosum hort. non
L., Vaccinium×covilleanum But. et Pl.) is relatively a new fruit crop when compared
to other fruit species. The first valuable cultivars (“seven big”) were introduced
in 1950s. It is a hybrid of at least three species (Vaccinium australe Small, V.
corymbosum L., and V. angustifolium Ait.) (1). The Vaccinium genus also contains
other useful species among others: cranberry (Vaccinium macrocarpon Ait.),
lingonberry (Vaccinium vitis-idaea L.), and bilberry (Vaccinium myrtillus L.). The
commercial world production of blueberries is 214–331 thousand tons, grown in the
area 48.3–74.0 thousand hectares during the period 1999–2008 (FAOSTAT). The
analogous production of cranberries was 286–440
Maurizio Lambardi et al. (eds.), Protocols for Micropropagation of Selected
Economically-Important Horticultural Plants, Methods in Molecular Biology, vol.
994, DOI 10.1007/978-1-62703-074-8_5, © Springer Science+Business Media New York
2013
63
64 W. Litwin czuk
thousand tons on 20.2–22.6 thousand hectare area (ibid). Several species of
Vaccinium are highly valuable for their edible fruits, which are excellent source
of health-promoting nutrients, among others: dietary fibers, antioxidants (vitamin
C, beta-carotene, folic acid, ellagic acid, anthocyanins), and antibacterial and
antifungal agents, like anthocyanosides, benzoic acid. The berries are con- sumed
either as fresh or processed products such as frozen, canned, liquid, concentrates,
and dried goods, like preserves, candies, jells, juices, syrups, ice creams,
pastries, muesli, milk and alcohol bever- ages, as well as nutritional supplements
and pharmaceuticals. There is an increase in highbush berries demand and fetch
attractive price, and that has rapidly expanded its cultivation area worldwide in
the last decades of twentieth century. In order to meet the increasing demand of
blueberry plantlets, micropropagation would be the most appropriate approach. The
micropropagation is the only method for rapid propagation of elite breeding clones
and new cultivars. By the end of 1980s more than one million highbush blueberry
plants were propagated annually worldwide (2). It should be noted that at the same
time some controversies about value of micropropagated plants, among others of
Ericaceae family, appeared (3, 4) and, although rare, remain until now. They were
based on some sporadic but confusing mentions of excessive vegetative growth (too
many thin lateral shoots), delayed fruit harvest for some years, or smaller
berries. However, despite of it at the turn of the twentieth/twenty-first century,
micropropagated plants prevailed conventional ones in nurseries. The first
articles about micropropagation of Vaccinium sp. appeared at the turn of the
1970/1980s. Since then numerous studies regarding blueberry (Vaccinium sp.) in
vitro cultures have been published. In fact, both blueberries and cranberries are
micropropagated with similar procedure—subculture of shoot explants on the medium
supplemented with indole-3-acetic acid (IAA, 0–4 mg/L) and 6-γ,γ-
dimethylallylaminopurine (2iP, 5–10 mg/L). However, intra- and inter-species differ
greatly in the response of Vaccinium sp. cultivars to the type and concentration of
cytokinins, auxins, and Fe salts (see Fig. 1). Therefore, it is recommended to
optimize culture conditions for each genotype separately. Also, explants easily
form adventitious shoots, even during initiation stage of Vaccinium in vitro
cultures (5–9). As adventitious shoots are better habituated to in vitro conditions
than axillary shoots, they could rapidly (in the few passages) displace axillary
shoots from micropropagation procedure. Thus, the special care should be taken to
prevent such phenomena as adventitious shoots of many species are suspected to be
the main source of somaclonal variation. The in vitro culture techniques are also
utilized in cryopreservation (10), in vitro selection (11), mutagenesis (12),
interspecific and intersectional hybrids (13), genetic transformation (3, 14, 15),
and production of secondary metabolites of pharmaceutical value in suspension and
callus cultures (16, 17).
Fig. 1. Comparison of the growth of Vaccinium sp. in vitro cultures on the same
medium. (a) highbush blueberry ‘Bluecrop’,(b) highbush blueberry ‘EarlyBlue’, (c)
highbush blueberry ‘Blueray’, (d) highbush blueberry—false ‘Bluecrop’, (e) highbush
blueberry ‘Herbert’, (f) cranberry ‘Stevens’, (g) cranberry ‘Pilgrim’, (h)
Vaccinium microcarpon ‘nn’, (i) lingonberry ‘Runo Bielawskie’.
66 W. Litwin czuk
2. Materials
2.1. Surface Sterilization of Source Material
In this chapter, the protocol is described for axillary shoot initiation and
multiplication in vitro, with subsequent rooting shoots in vivo along with
acclimatization of regenerated plants.
1. Ethanol 70% (v:v).
2. Commercial bleach solution (e.g. “ACE” bleach; 5% (v:v) NaClO), diluted 1:5
(v:v) with tap water or solution of 0.1% mercuric chloride.
3. Distilled (or autoclaved reverse-osmosis) water (150 mL aliquots in 300 mL screw
capped jars).
4. Laboratory shaker or ultrasonic washer.
5. Potted parental plant as a source of explants.
6. Solutions of Bayleton 5 WP (1.0 g/L), Topsin M (1.5 g/L), or equivalent
fungicides to spray parental plants.
1. Labwares: autoclave, magnetic stirrer with heating, dry-heat sterilizer,
microwave oven, pH meter, refrigerator, horizontal laminar flow bench.
2. Instruments: scalpels, forceps, glass bead sterilizers, or gas burners.
3. Sterile Petri dishes (100 × 15 mm) or sterile paper sheets (keep at 160°C for
minimum 2 h) to prepare explants.
4. Sterile dispensers of liquid medium to make double-phase medium, like graduated
(25 mL) pipettes with pumps, autoclavable and adjustable (1–5 mL) single-channel
pipettes.
5. The media for (a, b) culture initiation, and (c, d, e) shoot proliferation (see
Tables 1 and 2).
6. 1 M HCl and 1 M NaOH.
7. Test tubes, 15–75 mL capacity, with caps (16 × 75, 25 × 150 mm) or 25–50 mL
conical Erlenmeyer’s flasks.
8. Aluminum packaging foil (0.04 mm thick) for culture initiation and first
subcultures.
9. 300–500 mL glass jars closed with autoclavable, transparent lids (or adequate
culture vessels) for shoot proliferation and rooting.
10. Plastic, thermostable (0.1–3 L) beakers.
11. Culture room with air conditioning. The possibility of temperature (22–28°C),
light intensity (30–50 μmol/m2/s PPFD), and photoperiod (16/8 hday/night)
regulation is advantageous. The white growth shelves with white fluorescent lamps,
preferentially with bottom-cooling system.
2.2. Tissue Culture Facilities and Culture Media
5 MicropropagationofVacciniumsp. 67
Table 1
Composition of media used for micropropagation of Vaccinium sp.
Stages
(a) Initiation (liquid)
(b) Initiation (solidified)
(c) Proliferation (solidified)
(d) Proliferation—last passage (solidified)
(e) Proliferation—last passage (liquid)
Macronutrients
10–50% BMa 50% MS
10–50% BM 50% MS
100% BM 100% MS
100% BM 100% MS
100% BM 100% MS
Micronutrientsb (without Fe salts)
FeNaEDTA + FeEDDHA Vitaminsc
Myo-inositol
Sucrose
36.7 + 5 mg/L
36.7 + 5 mg/L
58.7 + 10 mg/L 100% WPM
100 mg/L
30 g/L −/5–10/80 mg/L IAA or IBA
58.7 + 10 mg/L 100% WPM 100 mg/L 30–40 g/L −/5–10/80
58.7 + 10 mg/L 100% WPM 100 mg/L 30–40 g/L
Cytokininsd (ZEA/2iP/AS) Auxinsd
0.5–1/80 mg/L
0.5–1/80 mg/L
mg/L
– –
Agar (see Note 9) Other ingredients
–
5–6 g/L
0–1.0 mg/L
6–11 g/L Fe-EDDHA 10 mg/L
6–11 g/L Fe-EDDHA
–
Fe-EDDHA 10 mg/L
pH
5.0
5.0
5.0
5.0
5.0
100% WPM
100% WPM
100 mg/L
100 mg/L
15 g/L
15 g/L
IAA 1.0 mg/L or IBA 0.2 mg/L
IAA 1.0 mg/L or IBA 0.2 mg/L
–
fructose (5 g/L), (PVP 360, 100 mg/L), L-cysteine 5 mg/L PPMTM (0.5 mL/L),
fructose(5 g/L), (PVP 360, 100 mg/L), L-cysteine 5 mg/L PPMTM (0.5 mL/L),
10 mg/L
aBM—one of media recommended for micropropagation of Vaccinium sp (see Note 6)
b MS micronutrients: NaFeEDTA 36.7 mg/L, KI 0.83 mg/L, H3BO3 6.3 mg/L, MnSO4 × 4H2O
22.3 mg/L, ZnSO4 × 0.25 mg/L, CuSO4 × 5H2O 0.025 mg/L, CoCl2 × 6H2O 0.025 mg/L
c Woody Plant Medium (WPM, (23)) vitamins: glycine 2.0 mg/L, thiamine HCl 1.0 mg/L,
pyridoxine HCl 0.5 mg/L, d Cytokinins, auxins (see Notes 8 and 11)
7H2O 8.6 mg/L, Na2MoO4 × 2H2O nicotinic acid 0.5 mg/L
68 W. Litwin czuk
Table 2
Composition of macronutrients in media used in in vitro cultures of Vaccinium sp.
Compounds (mg/L)
AND_M (20) AND_Z (21) EC (22) PMN (19) WPM (23) Z-2 (5) DR (24)NH4NO3
400
400
400
480
400
160
550
KNO3
190
480
202
305
–
202
330
K2SO4
–
–
–
–
990
–
140
(NH4)2SO4
–
–
132
–
–
198
50
Ca(NO3)2 × 4 H2O
684
–
–
470
556
708
410
CaCl2 × 2 H2O
–
440
440
–
–
–
200
KH2PO4
370
–
408
–
170
408
220
KH2PO4 × H2O
–
–
–
205
–
–
NaH2PO4 × H2O
–
380
–
180
–
–
100
MgSO4 × 7 H2O
370
370
370
370
370
370
370
Na2EDTA
74.6
37.2
–
73.2
74.6
74.4
FeSO4 × 7 H2O
55.6
27.9
–
60.0
55.6
55.7
NaFeEDTA––56–––37.32.3. Acclimation of Regenerated Plants to Ex Vitro Conditions
1. Greenhouse or plastic tunnel facilities or as a last resort well separated
culture room with air conditioning, provided with high pressure sodium lights (60–
200 μmol/m2/s PPFD).
2. Mist chambers (95% RH), multicell plug trays with transparent plastic
covers/lids. Cell dimensions about 2×3 cm (width, depth), transparent plastic foil.
3. Peat and sand mixture (2:1; v:v) adjusted to pH 4.0 with calcium carbonate,
watered with solution of SCOTTS Peters Plant Starter (10+52+10; 0.8 g/L) and
fungicide Previcur 607 SL (1.0 g/L) or equivalent chemicals.
4. Solution of Rovral Flo 255 SC (1.0 g/L) or equivalent fungicide to spray
microplants.
= 0.
(a) Find a solution of the D.E. and B.C. that satisfies the I.C. to within an error
of .00l.
(b) By computing Utt and Uxx at (x,t) = (0,0), show that there is no C 2 solution
of the problem.
7. Consider a string which vibrates in the xy-plane, but not necessarily
transversely. When the
string is at rest the points on the string have coordinates of the form (x,O) (0 ~
x ~ L). Suppose
that at time t, the point which was in the rest position (x,O), has xy--
<:oordinates (r(x,t),u(x,t))
[e.g., for transverse vibrations r(x,t) = x and for longitudinal vibrations u(x,t)
= 0].
(a) Show that the local stretching factor s at the point corresponding to x is
(r~ + u~)! .298
Chapter 5
The Wave Equation
(b) Suppose that the tension at the point corresponding to x is still of the form
g( s)T o. Show
that by virtue of Newton's equation ( F = rna) r(x,t) and u(x,t) satisfy the system
r tt
where s
= (r~ + u~)t,
[ To ~ ux] ,
with B.C. r(O,t) = u(O,t) = u(L,t) = ° and r(L,t) = 1.
= b~
[To ¥)r x]
and
u tt
= b~
(c) When will these equations decouple, in the sense that they can be solved
separately?
(d) The equations are still valid if D and To are allowed to be positive c1
functions of x, but
then To(x) cannot be brought outside of the parentheses. Assuming that g(s) = s,
find the
unique time-independent (steady-state) solution (R(x),U(x)) of the system with the
given B.C.,
f ° [To(x)]-l dx <
standard configuration (x,O), ° x
assuming that
L
~
00.
~
When is R(x):: x?
(Le., when the string is in the
L, under what circumstances is the string really at rest ?)
8. For a string that vibrates transversally in a medium, say air, one must take air
resistance into
account. Assuming that the force due to the air resistance is proportional (but
oppositely
directed) to the velocity ut(x,t) j, show that u(x,t) obeys the equation Utt = a
2uxx - kU t ' for
some real k> 0.
9. Use separation of variables to find all product solutions of the problem (with k
> 0)
D.E. Utt
= a 2 Uxx - ku t ,
B.C. u(O,t)
= 0,
u(L,t)
° x
~
= 0,
for the string with air resistance and fixed ends.
~
L,
-00
< t < 00,Section 5.2
299
D'Alemberl's Solution for Wave Problems
5.2 D'Alembert's Solution for Wave Problems
We have seen that one can represent the solution of the problem (where N < 00)
2
= a uxx '
0 ~ x ~ L, -w < t < 00,
B.C. u(O,t) = 0 u(L,t) = 0 ,
D.E.
I.C.
U tt
u(x,O)
= f(x) =
N
2n=lBnsin(r), ut(x,O)
N
(1)
= g(x) = 2n=lAnsin(r)
by the series
(2)
If f(x) and g(x) are not finite Fourier sine series, but are continuous and
piecewise C 1 and
vanish at x = 0 and x = L, we may approximate f(x) and g(x) to within any
(positive)
experimental error by truncations of their Fourier sine series (cf. Theorem 1 of
Section 4.3). Thus,
problem (1) has been solved for all practical purposes, for such f(x) and g(x), as
nearly as
anyone can say. However, in order that the theory have predictive value, we need to
know that
small changes in f(x) and g(x) induce small changes in the solution. Otherwise, two
different
approximations, both within experimental error, may lead to significantly different
solutions.
This property of "continuity of solutions with respect to variations in the I.C."
was established
for the heat equation by the use of the Maximum Principle (cf. Theorem 2 of Section
3.2).
However, a direct translation of the Maximum Principle to the case of the wave
problem (1) is
false, as the following example shows.
Example 1. In problem (1) take f(x) = 0 and g(x) = sin(1lX/L). Show that the
maximum of the
solution u(x,t) does not occur when t = 0, x = 0 or x = L.
Solution. The solution is u(x,t) = ;asin(7rat/L)sin(1lX/L). Note that u(x,t)
vanishes at the
ends and also initially (Le., u(O,t) = 0, u(L,t) = 0, u(x,O) = 0). However, u(x,t)
¢ 0 as a direct
translation of the Maximum Principle for the heat equation would imply. Indeed, the
maximum
of u(x,t) occurs at x = tL, t = tL/a (as well as t = (2n + t)L/a, n = 0, ±1, ... ).
0
The reason for the failure of the direct translation of the Maximum Principle is
that it does
not take into account both of the I.C. in (1). Eventually (cf. Theorem 5), we prove
that the
solution u(x,t) of (1) obeys the following type of maximum principle:
max lu(x,t)1
O~x~L
-w<t<oo
~ max If(x) I +~ max Ig(x)l.
O~x~L
a O~x~L
(3)300
Chapter 5
The Wave Equation
Note that this involves both f(x) and g(x), and the absolute values cannot be
removed. Using
(3), we will establish (cf. Corollary of Theorem 5) the desired result that small
changes in the I.C.
produce small changes in solutions. This was done for the heat equation in Theorem
3 of Section
3.2. The key to obtaining (3) is D'Alembert's formula for the solution of the wave
equation on
an infinite string (-00 < x < (0) with I.C. u(x,O) = f(x) and ut(x,O) = g(x) ,
namely,
1
1 JX+~
u(x,t) = 2[f(x-at) + f(x+at)] + 2aa
g(r) dr .
(4)
x-at
This formula is of great interest in itself, and it avoids the problem of
convergence of infinite series
in the Fourier series approach. Our first goal is to derive D'Alembert's formula
(4).
Derivation of D'Alembert's formula
We can write the wave equation in the form
[:2 - a 2 ~]U(X,t) = 0,
(5)
where the expression in parentheses is a differential operator which operates on
the function u to
yield U tt - a 2 u xx . This operator can be factored into two first-order
operators:
[fi2
at 2 -
2
rP ]
a &2 u =
[a at - a ox
a ] [a at + a ox
a ] u .
(6)
We can use this factorization to find the general solution of the wave equation.
u(x,t) is any C 2 solution of (5). Note that
Suppose that
In other words, the function y(x,t) == ut + au x solves the PDE Yt - ayx = 0. Since
the
characteristic lines (cf. Section 2.1) of this PDE are of the form x+at = const.,
we know that the
solution y must be of the form y(x,t) = h(x+at) for some
u t + aux
c1
= y(x,t) = h(x+at)
function h. Thus,
.
The characteristic lines for this first-order PDE for u are x-at
the right-hand side h(x+at), we make the change of variables
= const..
In view of the form ofSection 5.2
301
D'Alembert's Solution for Wave Problems
w = x - at
and
z = x + at .
Letting v(w,z) = u(x,t), we obtain
ut + au x
Thus, v(w,z)
=
I
= vww t
+ vzz t + a(vwwx + vzz x )
ia h(z) dz + G(w)
=
= 2avz = h(z)
.
F(z) + G(w), or
(7)
u(x,t) = F(x+at) + G(x-at) .
Hence, we have shown that an arbitrary solution of the wave equation can be written
in the form
(7), where F and G are arbitrary C2 functions. Converselx, one can easily check
that any
function of the form (7) is a solution of the wave equation (Le., t 7) is the
general solution). If we
graph the function F(x+at) as a function of x at a fixed time t, we obtain the
graph of F(x)
translated to the left by a distance of "at", as the reader may verify. Thus,
F(x+at) describes
a wave with initial profile F(x) moving to the left with speed a. Similarly, G(x-
at) yields a
wave traveling to the right with speed a. We have shown that the general solution
of
U tt = a 2 u xx is a superposition of two waves traveling in opposite directions
with speed a.
Example 2. We know that cos(Aat)sin(Ax) is a (product) solution of U tt = a 2u xx '
Hence, it
must be possible to rewrite cos(Aat)sin(Ax) in the form F(x+at) + G(x-at). Do it.
Solution. Using the identity cos(,8)sin( a)
cos(Aat)sin(Ax) =
= HSin( a+,B)
+ sin( a-,B)], with a
= Ax
and ,8 = Aat,
~ [sin(A(x+at)) + sin(A(x-at)] .
This exhibits the "standing wave" product solution, as a superposition of waves
traveling to the
right and left with speed a. 0
Theorem 1 (D'Alembert's Formula). Let f(x) be C2 and let g(x) be C 1
the unique solution of the problem
D.E.
U tt
2
= a uxx '
I.C. u(x,O)
= f(x),
-00
ut(x,O)
< X < 00,
-00
(-00
< x < 00). Then
< t < 00,
= g(x),
(8)
is given by
u(x,t)
= 2 1 [f(x+at) + f(x-at)]
1 Ix+at
+ -2
g(r) dr .
a x-at
(9)Chapter 5
302
The Wave Equation
Proof. We know that if a solution of (8) exists, then it must be of the form u(x,t)
= F(x+at)
+ G(x-at), where F and G are C2 functions. The I.C. will be satisfied precisely
when
f(x) = u(x,O) = F(x + O·a) + G(x - O·a) = F(x) + G(x), (10)
g(x) = ut(x,O) = F' (x)a - G' (x)a. (11)
Integrating the second equation, we obtain the following pair of equations for the
unknown
functions F(x) and G(x) :
F(x) + G(x)
= f(x)
and
F(x) - G(x)
=
if:
g(r) dr + C,
where C is an arbitrary constant. Adding and subtracting yields
F(x) = Hf(x) +
and
G(x)
= Hf(x) -
U:
U:
(12)
g(r)dr + C],
g(r)dr -
c] = Hf(x) + U: g(r)dr - c].
(13)
These equations are identities in the sense that they hold for all values of x, as
in sin2(x)
= 1 - cos 2(x). Because of this, we may substitute x + at for x in (12) and x - at
for x in
(13), and obtain valid results [e.g., sin 2(x+at) = 1 - cos 2(x+at)], namely,
°
1fxtat g(r)dr + C ] ,
F(x+at) = 2" 1 [ f(x+at) + a
G(x-at) = -21 [f(x-at) +
lfO
g(r)dr - C].
a x-at
(15)
Adding these expressions, we obtain (9). However, the above argument was based on
the
assumption that a solution of the problem (8) exists. We have just shown that if a
solution
exists, then it must be given by the D'Alembert's formula (9). We must finally show
that u(x,t),
given by (9), actually solves problem (8). Note that F(x) and G(x) defined by (12)
and (13)
are C2, since f(x) is C2 and g(x) is C1 ((g(r) dr is C2, since its derivative g(x)
is cl).
°
The right-hand side of (9) is F(x+at) + G(x-at) for the C2 functions F and G, and
hence we
know that u(x,t) defined by (9) solves the D.E.. The initial conditions of (8) are
met, since F(x)
and G(x) given by (12) and (13) satisfy (10) and (11), by construction. 0
°
Remark. There are no boundary conditions in problem (8), because the string has no
ends. For a
finite string, say
$ x $ L, the functions f(x) and g(x) would only be defined on [O,L], and
f(x±at) would be undefined for t l
UNITATE
C
Ă DE BAZĂ
Ă A EVIDE
ENȚEI EC ONOMICE
E
1. Obiective e
1.1

a

C
Contabilitat
s
mult mai
p
dub blă într-o
metrie. Dup
A
Astăzi,
con
c mai comp
o
într-un m
Princip
r reţea de
(după N.
N Feleagă) )
indiferent de
reprezentarea unei
în
economici,
rezultatele
financiare finale.
organizeze şi
Organizarea
dar şi o
organizatoric,
financiar-
contabilă.
cadrul
contabil. În
superioare),
superioare
loc
calendaristic,
calcula diverşi
pozitivi şi
contabilitate
patrimoniului;
lor.

entităţi.

unei entităţi.
 Ce este contabilitatea financiară şi prin ce se c
Utilizarea feromonilor
Dintre exohormonii cunoscuţi la insecte cea mai importantă grupă o reprezintă
feromonii, iar dintre aceştia interes practic deosebit prezintă atractanţii
sexuali. Ei se
folosesc în protecţia culturilor prin două căi: prin distrugerea în masă a
masculilor cu
ajutorul capcanelor cu atractanţi sexuali şi prin dezorientarea masculilor prin
perturbarea transmisiei feromonale normale.
Metoda capturării în masă a masculilor. Principiul combaterii se bazează pe
competiţia dintre atractantul sexual sintetic specific sub formă de capsule din
capcane
(fig. 58), prezent în doze corespunzătoare, cu atractantul sexual natural al
femelelor,
degajat în cantităţi infime. Din cauza acestei competiţii masculii se vor orienta
către
capcane şi vor fi distruşi, împiedicându-se astfel fecundarea femelelor din
populaţie.
Distrugerea masculilor în capcane se poate face prin diferite procedee. Metoda cea
mai
obişnuită constă în ungerea peretelui intern al părţii inferioare a capcanelor cu o
substanţă adezivă, care imobilizează masculii ce vor veni în contact cu ea. Numărul
capcanelor feromonale variază după specii între 5 - 40. Atractanţii sexuali cel mai
mult
experimentaţi în ţara noastră prin această metodă sunt: AtraPOM, AtraFUN,
AtraBLANC, AtraBOT, AtraBRAS. Rezultatele experimentale efectuate până în
prezent sunt promiţătoare (G h i z d a v u, 1982, 1983, 1984; G h i z d a v u şi
colab.,
1987 a, b; F i l i p, 1988; Z a h a r i a, 1998; N i c o l e s c u M i h a e l a -
A l e x a n d
r a, 1998).
Cele mai bune rezultate se pot obţine cu ajutorul feromonilor sexuali care atrag
ambele sexe sau femelele, dar numărul speciilor de insecte la care se folosesc
astfel de
atractanţi este foarte mic.
Metoda dezorientării masculilor sau a perturbării transmisiei feromonale
normale. Combaterea dăunătorilor prin această metodă se bazează pe principiul
reducerii potenţialului de reproducere a insectelor prin împiedicarea întâlnirii
celor două
sexe în vederea împerecherii. În acest scop se cunosc în prezent şi se pot folosi 7
tipuri
de produse dezorientante sintetice, dintre care enumerăm pe cele mai importante:
feromonii (atractanţii) sexuali, paraferomonii (atractanţi care nu sunt feromoni
specifici,
dar induc acelaşi răspuns comportamental) şi antiferomonii sau inhibitorii sexuali.
Astfel, dacă în timpul împerecherii atmosfera ecosistemului supus tratamentului
este
impregnată uniform cu o doză suficientă de atractant sexual sintetic sau de alt
produs
dezorientant, masculii sunt "dezorientaţi", nu mai sunt în stare să găsească
femelele,
care rămân nefecundate şi depun ouă nefertile. În acest mod efectivul şi densitatea
populaţiei dăunătoare scad simţitor. În ţara noastră s-a experimentat tehnica
evaporatoarelor distanţate, instalate manual. Dozele de atractanţi sexuali
folosite, diferit
după specie, sunt cuprinse între 2 - 40 g/ha.
În ţara noastră metoda dezorientării masculilor cu ajutorul atractantului sexual
sintetic specific a fost cercetată pentru combaterea viermelui merelor (G h i z d a
v u
şi colab., 1987 a) şi speciilor de buha verzei şi buha grădinilor de legume (N i c
o l e s c
u M i h a e l a - A l e x a n d r a, 1998).
117Fig. 58 - Capcane feromonale folosite în capturarea insectelor:
a - tip "Delta"; b - tip "Hexatrap"; c - tip "Tetratrap"; d - stativ
pentru capcane amplasate în culturi de legume (după G h i z d a v u).
Unele cercetări cu privire la dezorientarea masculilor prin folosirea inhibitorilor
sexuali au fost efectuate numai la buha verzei (S t a n şi colab., 1996; N i c o l
e s c u
M i h a e l a - A l e x a n d r a, 1998) şi buha semănăturilor (S t a n şi colab.,
1996).
Metodele biotehnice pot fi folosite cu succes în combaterea directă a unor specii
de lepidoptere noctuide dăunătoare, constituind una din verigile importante ale
combaterii integrate.
REZUMAT
Metodele folosite în combaterea dăunătorilor animali ai plantelor cultivate, după
scopul lor se clasifică în preventive şi curative. Metodele preventive şi curative
de
combatere a dăunătorilor animali, după mijloacele folosite, se grupează în
următoarele
categorii: măsuri de carantină fitosanitară, metode agrofitotehnice, folosirea de
hibrizi şi
soiuri de plante rezistente, metode mecanice, metode fizice, metode chimice şi
metode
biologice.
In general metodele agrofitotehnice au un caracter profilactic, prin aplicarea lor
realizându-se în primul rând, prevenirea atacului dăunătorilor precum şi reducerea
înmulţirii lor şi a atacului.
118Pentru obţinerea unor eficacităţi ridicate, metodele agrofitotehnice trebuie să
fie
utilizate pe perioade lungi, cu mult înainte ca paguba să devină vizibilă.
In ceea ce priveşte utilizarea de soiuri sau hibrizi rezistenţi, aceasta reprezintă
o
metodă ideală, deoarece implică un cost minim, nu poluează mediul înconjurător şi
nu
afectează fauna utilă.
Combaterea dăunătorilor cu substanţe chimice s-a dovedit, până în prezent,
foarte eficace, substanţele utilizate având o viteză de acţiune ridicată, forme
variate de
condiţionare şi posibilitatea tratării unor suprafeţe mari.
Extinderea metodelor biologice de combatere a dăunătorilor este una dintre
direcţiile de perspectivă a dezvoltării protecţiei plantelor, aceste metode ocupând
un loc
însemnat în practica agricolă. Lupta biologică este rezultatul unei intervenţii
active a
omului şi constă în folosirea diferitelor mijloace şi metode biologice în
distrugerea
dăunătorilor animali în scopul reducerii densităţii numerice a populaţiilor
acestora sub
pragul economic de dăunare.
ÎNTREBĂRI
8.1. Ce este carantina fitosanitară?
8.2. Care este rolul asolamentului?
8.3. Ce înţelegeţi prin plantă rezistentă?
8.4. Prezentaţi pe scurt principalele metode mecanice.
8.5. Care sunt avantajele utilizării metodelor chimice? Dar dezavantajele?
8.6. Ce este agrodisponibilitatea?
8.7. Ce este selectivitatea?
8.8. Care sunt principalele tipuri de selectivitate?
BIBLIOGRAFIE
8.1. Alexandrescu Sanda, Baicu T. (1973) - Utilizarea inhibitorilor de nutriţie în
protejarea culturii de cartof contra gândacului din Colorado (Leptinotarsa
decemlineata Say), Anal. I.C.P.P., vol. IX, p. 449 - 456.
8.2. Alexandrescu Sanda, Prunescu C., Traciuc Elena, Codreanu Doina (1975) -
Modul de acţiune al Brestanului şi oxiclorurii de cupru ca inhibitori de nutriţie
ai
gândacului din Colorado (Leptinotarsa decemlineata Say), Anal. I.C.P.P., vol.
IX, p. 231 - 245.
8.3. Baicu T., Săvescu A. (1978) - Combaterea integrată în protecţia plantelor,
Edit. Ceres, Bucureşti.
8.4. Bonnemaison L. (1961) - Les ennemis animaux des plantes cultivées et des
forets, vol. I, Paris.
8.5. Călin Maria (1996) - Cercetări cu privire la biologia şi combaterea integrată
a dăunătorilor la varză, Teză de doctorat, U.S.A.M.V., Bucureşti.
8.6. Ciochia V. (1982) - Tehnologia creşterii semiindustriale a entomofagului
Trichogramma Westw. în vederea utilizării acestuia în limitarea populaţiilor
unor dăunători ai sfeclei de zahăr din România (I), Lucr. Şt. Sfeclă de zahăr, XI,
Fundulea - Bucureşti, p. 163 - 182.
8.7. Ciochia V. (coordonator) şi colab. (1997) - Limitarea populaţiilor de
dăunători vegetali şi animali din culturile agricole prin mijloace biologice şi
biotehnice în vederea protejării mediului înconjurător. Edit. Disz Tipo, Braşov.
8.8. Ciurhii M. (1997) - Combaterea microbiologică cu bacterii şi ciuperci
entomopatogene a unor dăunători, în: "Limitarea populaţiilor de dăunători
vegetali şi animali din culturile agricole prin mijloace biologice şi biotehnice în
vederea protejării mediului înconjurător". Edit. Disz Tipo, Braşov, p. 201 - 257.
1198.9. Isac Gr. (1973) - Cercetări privind combaterea biologică a viermelui
merelor
(Carpocapsa pomonella L.) prin folosirea entomofagului Trichogramma
embryophagum Htg., Anal. I.C.P.P., vol. IX, p. 377 - 391.
8.10. Knipling E. F. (1979) - The basic principles of insect population
suppression and management, U.S. Dept. Agric. Handbook 512, 659 p.
8.11. North D. T (1975) - Inherited sterility in Lepidoptera, Ann. rev. Entomol.,
20, p. 167 - 182.
8.12. Şandru Gavrilă (1992) - Cercetări privind controlul biologic al unor
dăunători din culturile agricole cu ajutorul speciilor de Trichogramma în judeţul
Mureş, Teză de doctorat, U.S.A.M.V., Bucureşti, 186 p.
1209. DĂUNĂTORI POLIFAGI
CUVINTE CHEIE: lăcuste, gândacii pocnitori, viermi albi.
OBIECTIVE
- prezentarea principalelor specii de dăunaracterizează.
13arge, making the solution (9) undefined. We overcome this
problem later by extending (in various ways that depend on the B.C.) f(x) and g(x)
toen-
tration of the second messeAceastă închidere istorică a marcat evoluţia statului
român în lumea
modernă, evoluţie care a putut fi reorientată numai după instaurarea ordinii
burgheze propusă de revoluţiile de la 1848 care vizau, în principal,
sincronizarea României cu Occidentul dezvoltat.
2.1 Evoluţia satului românesc
în perioada formării burgheziei româneşti
Până la începutul secolului al XIX- lea, societatea românească era
preponderent rurală. La stabilirea acestei preponderenţe a contribuit
situarea Ţărilor Româneşti la intersecţia imperiilor, presiunile exercitate de
acestea făcând ca, din secolele XV- XVI, atracţia pentru o viaţă urbană să
se diminueze întrucât comerţul era strict limitat, iar industriile se reduceau
la satisfacerea necesităţilor stricte ce ţineau de economia naturală.
Din această cauză, la începutul secolului al XX-lea, cea mai mare
parte a populaţiei era stabilită, în continuare, în spaţiul rural. Astfel, la o
populaţie de 7,5 milioane de locuitori ai vechiului regat în 1914, peste 90%
erau locuitori ai aşezărilor rurale. În anul 1914, însă, România se găsea
deja pe calea modernizării, începută sub impactul ideilor liberale vehiculate
de revoluţiile de la 1848. Desigur, revoluţiile de la 1848 nu au izbutit să
realizeze cadrul obiectiv al unor transformări radicale, cadru ce a
devenitSociologie economică rurală
însă posibil abia după Unire şi după eliberarea de stăpânirea turcească.
Constituţia liberală din 1886 stabilea cadrul legal, întărit de Codul liberal
adoptat la 1886.
Funcţiunea unei economii de piaţă în România nu era însă posibilă
fără o infuzie de capital extern, întrucât statele româneşti nu reuşiseră să
realizeze acea acumulare primitivă care permitea dezvoltarea capitalului
comercial şi apoi a celui industrial.
O dată cu Reforma Învăţământului din 1864, când s-a stabilit
învăţământul general şi obligatoriu de patru ani, s-au creat premisele unei
valorificări a resurselor umane superioare pe teritoriul Principatelor.
Realizarea unei economii moderne, de tip capitalist, a fost
îngreunată de gradul scăzut al culturii de masă, comunitatea sătească
menţinându-şi o cultură tradiţională specifică, dar legată de zonele ei de
fixare. Cultura de masă presupunea o anumită uniformizare şi omogenizare
a tradiţiilor şi obiceiurilor care să facă posibilă circulaţia rapidă a ideilor şi
bunurilor. Economiile comunitare săteşti româneşti erau, până în preajma
anilor 1850, economii naturale închise, de subzistenţă, care, datorită slabei
dotări cu mijloace tehnice, nu asigurau acel surplus susceptibil de
valorificare pe piaţă.
Societatea românească era marcată şi de discrepanţele majore
dintre volumele proprietăţilor agricole, accentuate după aplicarea, începând
cu anul 1830, a Regulamentului Organic. În condiţiile în care produsele
occidentale pătrund în România, după revenirea la domniile pământene, se
constată o tendinţă a păturii marilor proprietari, a boierimii, de exploatare a
domeniilor prin intermediul arendaşilor. Arendaşii români nu erau însă
arendaşi în sensul occidental al termenului. În Occident, domeniile de
exploatare erau încredinţate pe termen lung unor manageri. Arendaşii din
România erau, în fapt, nişte interpuşi, pe termen scurt, pentru sume
forfetare, între proprietar şi domeniul exploatat, arendaşul urmărind să
obţină sume cât mai mari pe baza muncii ţăranilor, cărora li se impuneau
învoielile agricole.
Învoielile agricole au fost transformate prin Regulamentul Organic în
muncă prestată pentru proprietarii domeniilor, aşa-numita dijmă la tarla, şi
prelucrarea unei părţi a produselor de pe lotul propriu – dijma. Apariţia
arendaşilor transformă clasa proprietarilor într-o clasă de uzufructuari, iar
pe ţăranii învoiţi în adevăraţi robi, întrucât obţinerea profitului maxim
impunea creşterea continuă a dijmelor.
Premisele modernizării economiei româneşti, modernizare ce a
afectat iremediabil satul, sunt de două tipuri:
− externe, în contextul creşterii interesului Occidentului pentru
comerţul pe Dunăre şi Marea Neagră;
− interne, în contextul dispariţiei boierimii, ca urmare a apariţiei
economiei băneşti.Evoluţia istorică a comunităţii rurale din România
Războiul din Crimeea, provocat de Anglia şi sprijinit de Franţa şi
Sardinia din necesitatea de a exclude accesul Rusiei la Strâmtori, a impus
ca, la Convenţia de la Paris din 1858, să se stabilească nu numai posibila
unire a Principatelor, ci şi o serie de adaptări legislative care să le asigure
deschiderea spre comerţul cu Occidentul.
Principatele puteau fi considerate, desigur, un fel de colonii agricole
occidentale întrucât trebuia ca ele să-şi finanţeze importurile din ţările
europene dezvoltate prin exportul celor mai la îndemână resurse, în special
agricole. De aceea, din nevoia stimulării producţiei agricole, în special a
celei cerealiere, s-a realizat o reformă a proprietăţilor agrare, începută prin
secularizarea averilor mânăstireşti – 1863 – şi continuată cu reforma
propriu-zisă – 1864.
Creşterea producţiei cerealiere din necesităţi de export a implicat şi
o creştere a circulaţiei băneşti, ca mijloc de schimb, circulaţie convertită în
capital de camătă şi capital comercial. În acest sens, gânditorul german
W. Sombart, arată că tipurile de cămătărie apar, de obicei, în momentul în
care o economie, organizată până atunci pentru trebuinţele proprii, este
silită, din motive externe, să se prefacă în economie bănească.
Capitalul cămătăresc necesar economiei băneşti a fost realizat în
Occident, în special de populaţiile evreieşti, întrucât biserica socotea
camăta un păcat capital. S-ar putea spune, din această perspectivă, că
evreii au întemeiat capitalismul, întrucât ei realizau, prin secolele XV – XVI,
majoritatea operaţiunilor de creditare, capitalizare şi transfer bănesc. Până
în jurul anilor 1830, în România a funcţionat economia naturală
autosuficientă, care nu avea nevoie de agenţi specializaţi pentru circulaţia
banilor. O dată cu presiunile Occidentului pentru deschiderea pieţei
Principatelor şi, concomitent, cu apariţia ideilor liberale, nevoia de circulaţie
bănească impune apariţia acestor agenţi specializaţi, care au fost tot evreii.
Evreii au început să sosească în România din Polonia şi Ungaria,
având în spate credite de la comunităţile iudaice din zonă, pentru a
răspunde nevoilor circulaţiei băneşti impuse de producţia pentru piaţă.
Majoritatea dintre ei au practicat, iniţial, o formulă primitivă de capitalism,
prin camătă, dar această formulă a avut un rol distructiv major în raport cu
vechea proprietate funciară de tip boieresc, proprietate care, practic, era
desfiinţată după reformele liberale din 1864 şi 1866. Puterea boierească a
fost astfel înlocuită cu o oligarhie financiară liberală, preponderent
românească, întrucât evreii nu au fost naturalizaţi decât, relativ, târziu.
Oligarhia financiară impune fundamentarea materială a statului
modern, centralizat, dar şi apariţia mercantilismului, a spiritului de
economie, pe care majoritatea populaţiei cantonată în mediul rural nu îl
avea.
Transformarea producţiei de cereale într-o bază economică a
modernizat România, a impus crearea unor mijloace de comunicaţie
rapidă, de tipul căilor ferate şi şoselelor, crearea unor instituţii de credit
deSociologie economică rurală
tipul băncilor şi omogenizarea vieţii publice prin uniformizarea legislativă,
culturală şi lingvistică, realizându-se astfel modernizarea naţiunii române.
Regimul agrar al comunităţilor ţărăneşti a suferit o serie de transformări
care îl făceau accesibil adaptării la economia de piaţă şi la acumularea
bănească.
Până la revoluţiile liberale şi deschiderea spre importul occidental,
majoritatea ţăranilor români erau legaţi de pământ prin şerbie, rumânie sau
vecinie. În Transilvania, această legare a fost concretizată prin „Tripticul
Verboczi” din 1504, iar în Ţara Românească şi Moldova prin Regulamentul
Organic din 1829. În Transilvania, şerbia presupunea 52 de zile de robotă
pentru proprietarii terenurilor, în timp ce în Ţara Românească nartul sau
norma era de 56 de zile, iar în Moldova, de 84 de zile. În Basarabia, care a
fost răpită în 1812 de Imperiul Ţarist, în afară de dijma datorată
proprietarilor şi de dările către stat, ţăranii erau obligaţi să presteze 127 de
zile de muncă pe an. Deşi, atât în Moldova, răzeşii, cât şi în Muntenia,
moşnenii, au rezistat ca ţărani liberi, majoritatea ruralilor erau supuşi clăcii,
fiind obligaţi să împartă cu proprietarul produsele loturilor aflate în folosinţă.
Pătrunderea capitalismului occidental a implicat o serie de reforme
prin care loturile folosite au trecut în proprietatea ţăranilor. Astfel reforma
agrară din Transilvania a împroprietărit, prin despăgubire sau
răscumpărare, 140000 de familii. Această împroprietărire a fost confirmată
prin patentele imperiale din anii 1853 şi 1854. În Basarabia, ca urmare a
eliberării iobagilor din Rusia, s-a format, în anul 1869, o pătură de
proprietari incluşi în obşti, mărimea proprietăţilor fiind de la un ha, pentru
100000 de ţărani, la 3,5 ha pentru 200000 de ţărani şi peste 8 ha pentru un
număr de 15000 de ţărani. În cazul de împroprietărire prin răscumpărare, în
mod explicit, ţăranii erau siliţi să nu treacă la activităţi de tip urban şi să nu
părăsească mediul rural. Această restricţie se ridică abia în anul 1906,
după prima revoluţie burgheză care a zguduit Rusia.
În România, prin reforma de la 1864, au fost împroprietărite peste
486ooo de familii, care au primit din domeniile statului, dar şi din marile
latifundii, peste 1800000 ha. Această porţionare în mici proprietăţi a
domeniului agricol, necesară din punct de vedere social, pentru excluderea
presiunilor comunităţii rurale, nu a dus automat la creşterea producţiei
agricole, întrucât mijloacele şi cunoştinţele tehnice erau precare, iar
învoielile manipulate de arendaşi nu lăsau, practic, ţăranilor, nimic.
În aceste condiţii, statul a intervenit de mai multe ori pentru
modificarea, prin lege, a învoielilor, reducând termenul lor şi suspendând o
serie de dări aferente, ceea ce a permis creşterea producţiei, dar nu a dus
automat la îmbunătăţirea calitativă a vieţii rurale, ceea ce va provoca o
serie de răscoale cum au fost cele de la 1888 şi 1907, răscoale mistificate
ulterior din perspectiva numărului victimelor. Reprimarea acestora a fost
totuşi crâncenă, întrucât tânărul stat românesc, prins între două mari şiEvoluţia
istorică a comunităţii rurale din România
puternice imperii, austro-ungar şi rusesc, nu îşi putea permite să furnizeze
acestora pretextul unor intervenţii datorate dezordinii.
Trecerea de la capitalul cămătăresc la capitalul comercial al
oligarhiei financiare şi apoi la capitalul industrial a determinat presiunea
exercitată economic asupra societăţii, pentru adaptarea la vremurile noi,
dar abia prin reforma agrară de după primul război mondial s-a realizat o
redresare a satului românesc în faza propriu-zisă a modernităţii.
2.2 Satul românesc în epoca modernă
Epoca modernă a satului românesc începe, în mod paradoxal, în
ciuda Regulamentului Organic care a dus la legarea ţăranului de pământ în
1830, după tratatul de la Adrianopole. Prin acest tratat, Ţările Române,
care fuseseră debranşate de la comerţul european de ocupaţia turcească,
reintră în circuit, încep să participe la schimburile care se generalizează în
Europa. Principala beneficiară a Tratatului de la Adrianopole este Anglia, al
cărei interes impunea deschiderea comerţului pe Dunăre, axă majoră a
Europei şi stimulent natural în vederea satisfacerii nevoilor ei. În Ţările
Române, cu excepţia Transilvaniei încorporată în Imperiul Austro-Ungar,
Anglia era interesată, în primul rând, de produsele cerealiere. Agricultura
românească este astfel stimulată să producă pentru export, în primul rând
grâne necesare comerţului cu englezii.
Această stimulare impune o anume modernizare a agriculturii,
inclusiv a infrastructurii, prin care să se asigure transportul rapid al
produselor agricole în porturile de îmbarcare.
Dezvoltarea comerţului cu grâne are consecinţe majore asupra
satului românesc, deoarece schimbul nu era unilateral, cereale contra bani,
ci presupunea şi importul, atât a unor mijloace tehnice pentru creşterea
producţiei, cât şi a unor bunuri de consum. Pătrunderea capitalismului în
Ţările Române, ca urmare a comerţului, generează ceea ce Marx a numit
corect “despărţirea agriculturii de industrie”, sugerând formarea unei pieţe
interne pe care să se producă schimbul. Termenul de industrie este folosit
în sensul industriei casnice, majoritatea gospodăriilor rurale, în România
acelei perioade, fiind axate pe o economie de subzistenţă, după ce erau
satisfăcute nevoile posesorilor terenurilor.
Deschiderea spre comerţ, realizată ca urmare a necesităţii
exportului de cereale, va distruge industria casnică rurală, majoritatea
gospodăriilor ţărăneşti fiind după perioada muncilor agricole propriu-zise
mici ateliere în care se satisfăceau nevoi, cum ar fi: îmbrăcămintea,
repararea sau producerea de unelte agricole.
În lucrarea “Burghezia română”, Ştefan Zeletin observă consecinţele
acestei pătrunderi a capitalului într-o zonă blocată până atunci de
extinderea Imperiului Turcesc: “În România, factorul ce ruinase vechileSociologie
economică rurală
meserii era industria străină, nu indigenă şi, astfel, nevoile pieţei abia
născute trebuiau satisfăcute de străini, cu mărfuri aduse de peste hotare”. 3
De aceea, după nimicirea industriei mici, singurul factor producător de
valori proprii a rămas agricultura. Orăşenimea nu era producătoare
industrială şi cei care erau prinşi în mecanismul vieţii economice făceau
comerţ, adică asigurau circulaţia. Dar aceasta nu produce valori proprii, ci
doar pune în mişcare valorile produse de alţii. Şi astfel munca ţăranului a
rămas, la început, singurul izvor de venit în tânăra noastră societate
burgheză. Din ea, trebuia să trăiască ţăranul român, să fie întreţinută pe
calea bugetului orăşenimea, din ea trebuia să se plătească mărfurile aduse
din străinătate, ca şi imensele împrumuturi contractate de acolo.
Evoluţia ruralităţii, sub impactul cerinţelor pe piaţa externă de
produse agricole, a imprimat şi o anume restructurare a culturilor, cerinţele
fiind, în primul rând, de grâne, deşi ţăranul român îşi satisfăcea nevoile cu
porumb. Capitalul străin, realizat în urma exportului, era convertit în
produse tehnice de stimulare a agriculturii într-o proporţie relativ mică,
restul fiind cheltuit pe produse de lux de către marii proprietari. În aceste
condiţii, reforma agrară realizată de Cuza, ce a dus la împroprietărirea unui
mare număr de familii ţărăneşti, a permis o convertire a veniturilor din
agricultură către modernizarea acesteia. Această modernizare, impusă de
cerinţele din ce în ce mai mari de produse agricole, pe de o parte, nu a dus,
la refacerea industriei casnice rurale şi a impus, pe de altă parte, o tendinţă
de concentrare a proprietăţii agricole, concentrarea făcând ca cei care
dispuneau de venituri mai puţine să se proletarizeze, ajungâ
Evoluţia filogenetică se referă la etapele evoluţiei plantelor de-a lungul

generaţiilor şi este
influenţată de succesiunea condiţiilor de mediu din timpul fiecărei generaţii. Dacă
condiţiile de
mediu în care plantele s-au format se schimbă, apare un nou mod de manifestare,
apar însuşiri şi
caractere noi.
Schimbări ale modului de manifestare se întâlnesc la unele soiuri de ceapă care s-
au format
în condiţii de zi lungă. Dacă acestea sunt cultivate în condiţii de zi scurtă,
plantele nu formează
bulb. Această manifestare este folosită pentru practică, pentru obţinerea cepei de
stufat, care se
cultivă toamna sau primăvara devreme, când zilele sunt scurte. Salata, spanacul,
cultivate în condiţii
de zi lungă, formează o rozetă de frunze şi trec rapid la stadiul de înflorire şi
fructificare. Dacă se
cultivă în perioadele cu zile scurte, plantele formează o rozetă foarte bogată de
frunze, nu înfloresc,
şi acest lucru se valorifică în practică pentru obţinerea masei vegetative (a
frunzelor) care se
consumă. Intervenţia omului este evidentă în schimbarea condiţiilor de mediu prin
tehnologia de
cultură, ameliorarea plantelor, extinderea ariei de cultivare etc., mai ales dacă
schimbarea
succesiunii condiţiilor de mediu apare cât mai aproape de începutul vieţii
organismului. Omul, prin
intervenţiile sale, pe măsura aprofundării cercetărilor de biologie moleculară şi
ingineriei genetice, a
făcut posibilă crearea de soiuri şi hibrizi cu calităţi şi însuşiri superioare, la
care se manifestă
fenomenul heterozis, în prezent în legumicultură acest fenomen întâlnindu-se la
circa 20 de specii
(tomate, castraveţi, varză, ceapă etc.).
3.3.2. Evoluţia ontogenetică
Evoluţia ontogenetică se referă la etapele de evoluţie a organismelor în cursul
unei
generaţii şi înregistrează 3 perioade de viaţă:
1. Perioada de sămânţă, care cuprinde 3 faze:
- faza embrionară – se desfăşoară din momentul fecundării până la maturarea
seminţelor.
Noile organisme sunt foarte sensibile în această fază, prezintă cea mai mare
plasticitate ecologică şi
sunt strâns legate de planta mamă.
- faza de repaus – durează din momentul în care seminţele devin mature din punct de
vedere fiziologic, până când se declanşează procesul de germinare al acestora. În
această fază
procesele biochimice sunt mult încetinite, de aceea unele condiţii de mediu mai
precare din
perioada de păstrare, nu afectează viabilitatea seminţelor. Cu cât temperatura din
perioada de
păstrare este mai scăzută (4 - 5°C) şi umiditatea relativă mai mică, cu atât
perioada de păstrare a
seminţelor este mai mare.
29În această fază, seminţele trec prin 2 subfaze de repaus: repausul profund şi
repausul forţat.
Repausul profund începe imediat după recoltare şi se menţine o anumită perioadă de
timp
în funcţie de specie. Asigurarea unor condiţii optime de germinare nu pot duce la
declanşarea
procesului de germinare, întrucât repausul este considerat o necesitate biologică,
iar germinarea
seminţelor este posibilă după satisfacerea repausului.
Repausul forţat urmează repausului profund şi apare datorită lipsei sau
insuficienţei apei
şi căldurii.
- faza de germinare – începe din momentul crăpării tegumentului seminal şi ţine
până la
formarea primei frunze adevărate. În această fază plantele manifestă cerinţe
deosebite faţă de
umiditate (apa având rol esenţial în hidratarea seminţelor şi plesnirea
tegumentului), temperatură
(trebuie atins cel puţin plafonul minim de temperatură de la care se declanşează
germinarea
seminţelor) şi oxigenul care este indispensabil, lipsa acestuia în substrat ducând
la putrezirea
seminţelor, asociat şi cu alte cauze (umiditate excesivă, temperatură scăzută
etc.). După răsărire
plantele sunt foarte sensibile, firave şi supuse unui proces de selecţie naturală
foarte riguros. În
această fază supravieţuiesc plantele viguroase care provin din seminţe sănătoase,
mari, cu substanţe
de rezervă suficiente pentru hrănirea embrionului.
2. Perioada de creştere vegetativă, care cuprinde 3 faze:
- faza de răsad – care durează de la apariţia primei frunze adevărate şi până la
începerea
depunerii substanţelor de rezervă în organele specializate. În această fază, hrana
sintetizată de
plante este folosită în exclusivitate pentru creşterile vegetative, iar raportul
între procesele de
asimilaţie şi dezasimilaţie este aproape unitar (se consumă atât cât se produce).
Pentru asigurarea
creşterii normale în această fază, un rol important îl joacă factorii de vegetaţie
(temperatura, lumina,
umiditatea, hrana şi solul sau substratul) care trebuie să atingă valori apropiate
de cerinţele speciei
respective.
- faza de acumulare a substanţelor de rezevă – începe din momentul în care
surplusul de
hrană sintetizat de plante, se depune în organe specializate. Acestea îşi modifică
forma,
dimensiunile şi devin părţi comestibile ale plantelor legumicole. Depunerea
substanţelor de rezervă
se face în frunze, acestea cresc în dimensiuni (salată, spanac, sfeclă de frunze,
ţelină pentru frunze,
basela etc.), în tulpini subterane (cartof), în tulpini aeriene (gulie), în fructe
(tomate, ardei, vinete,
bame, fasole etc.), în primordii de inflorescenţă (conopidă, broccoli), în muguri
(vărzoase) etc.
- faza de repaus – este specifică plantelor bienale, trienale şi multianuale şi
începe din
momentul în care metabolismul plantei se reduce foarte mult datorită factorilor de
mediu
nefavorabili (temperatură scăzută, zile scurte). În această fază frunzele şi
rădăcinile active mor, cu
excepţia plantelor multianuale la care rădăcinile rămân active, însă procesele
metabolice se
desfăşoară într-un ritm foarte scăzut. Faza de repaus este şi o metodă de adaptare
a plantelor perene
la condiţiile de mediu, specifice ţării noastre, perenitatea fiind asigurată de
către organele subterane
în care se depozitează substanţele de rezervă. Intrarea în repaus mai este
influenţată şi de
acumularea acidului abscisic (Burzo, 1992) şi este controlată genetic.
Ieşirea din repaus diferă cu specia şi este determinată de scăderea concentraţiei
acidului
abscisic în muguri, de temperatura scăzută care stimulează biosinteza
giberelinelor, fapt care duce
la întreruperea repausului.
3. Perioada de reproducere, care cuprinde 3 faze:
- faza de îmbobocire – care începe odată cu apariţia bobocilor florali şi se
încheie cu
maturarea celulelor sexuale (a gameţilor). Această fază se caracterizează prin
creşterea suprafeţei de
asimilaţie, schimbarea compoziţiei chimice a organelor vegetative, iar recoltarea
părţilor
comestibile trebuie să se facă înaintea apariţiei tulpinilor florifere.
30- faza de înflorire – durează din momentul maturării gameţilor până în momentul
fecundării şi coincide cu deschiderea florilor la unele specii legumicole, dar se
poate desfăşura şi
înaintea deschiderii florilor; se impune alegerea corectă a momentului hibridării
pentru obţinerea
unor rezultate bune.
- faza de fructificare – ţine din momentul fecundării şi până când seminţele devin
mature
din punct de vedere fiziologic şi independente de planta mamă. Aceasta este faza
care încheie ciclul
ontogenetic al plantelor dintr-o generaţie şi faza care începe noua generaţie.
Plantele mamă de la
speciile anuale, bienale şi trienale se epuizează, iar embrionii se fortifică şi
aceştia vor forma noua
generaţie. La speciile multianuale, în faza de maturare a seminţelor, în organele
specializate se
depun substanţele de rezervă asigurând perenitatea speciei.
Parcurgerea acestor perioade diferă cu specia, şi anume:
- la plantele legumicole anuale: faza embrionară - faza de repaus - faza de
germinaţie -
faza de răsad - faza de îmbobocire - faza de înflorire - faza de fructificare,
suprapunerea
fazelor fiind evidentă;
- la plantele legumicole bienale: faza embrionară - faza de repaus - faza de
germinaţie -
faza de răsad (plantele în primele faze) - faza acumulării substanţelor de rezervă
- faza
de repaus - faza de îmbobocire - faza de înflorire - faza de fructificare;
- la plantele legumicole multianuale se trece prin perioada de sămânţă şi a
creşterii
vegetative cu fazele aferente în primii doi ani, iar apoi în fiecare an planta
înfloreşte şi
fructifică normal.
3.4. Particularităţile creşterii şi dezvoltării
Creşterea este un proces cantitativ, ireversibil, care contribuie la mărirea
dimensiunilor
plantelor (creşterea în înălţime, apariţia lăstarilor, creşterea frunzelor etc.).
Acest proces este
controlat genetic, dar foate mult influenţat de condiţiile de cultură (hrană, apă,
căldură, lumină) şi
determină apariţia tulpinii, frunzelor şi rădăcinilor.
Dezvoltarea reprezintă un proces calitativ care conduce la apariţia organelor de
reproducere. Florile din punct de vedere morfologic pot fi hermafrodite (solanacee,
fasole, mazăre)
şi unisexuate (cucurbitaceele). Majoritatea plantelor prezintă organele de
reproducere bărbăteşti
(staminele) şi organele femeieşti (pistilele) în aceeaşi floare (plante cu flori
hermafrodite). Florile
unisexuate (au un singur sex) pot fi pe plante monoice (flori unisexuate pe aceeaşi
tulpină) sau pe
plante dioice (flori bărbăteşti pe o plantă şi flori femeieşti pe altă plantă).
Plante dioice sunt:
spanacul, sparanghelul etc.
Polenizarea florilor la plantele legumicole poate fi:
- autogamă (autopolenizare) – când polenizarea are loc cu polen de la aceeaşi
floare sau
de la alte flori de pe aceeaşi plantă (polen propriu). Se întâlneşte la tomate,
ardei, vinete,
fasole, mazăre etc.
- alogamă (încrucişată) – când polenizarea se face cu polen de la alte plante. Dacă
polenizarea se face cu ajutorul insectelor, procesul se numeşte polenizare alogamă
entomofilă şi se întâlneşte la varză, ceapă, sparanghel etc. Dacă polenizarea se
face cu
ajutorul vântului, polenizarea este numită alogamă anemofilă (la spanac, sfeclă,
lobodă).
Plantele legumicole prezintă o serie de particularităţi privind creşterea şi
dezvoltarea.
31La cucurbitacee întâlnim pe aceeaşi plantă atât flori bărbăteşti, cât şi
femeieşti, dar
proporţia este diferită. Astfel, la unele soiuri, pe tulpina principală şi pe
ramificaţiile de ordin
inferior, se află mai multe flori bărbăteşti, iar pe ramificaţiile de ordin
superior predomină florile
femeieşti. Florile femeieşti se recunosc prin forma ovarului care este asemănătoare
cu a fructului.
Pentru a obţine un număr mai mare de flori femele, deci un potenţial productiv mai
mare, se
efectuează ciupiri repetate care stimulează ramificarea. De asemenea, fructele apar
şi cresc
concomitent cu formarea de noi flori, frunze şi cu creşterea plantei. Există soiuri
care formează
fructe fără fecundare, procesul fiind denumit partenocarpie, iar fructele
partenocarpice. De
asemenea, există hibrizi care formează, atât pe tulpina principală, cât şi pe
lăstari, numai flori
femele, sunt foarte productivi şi se numesc hibrizi ginoici. La hibrizii ginoici nu
este necesară
ciupirea repetată a plantei pentru stimularea ramificării şi formarea florilor
femele, făcându-se astfel
economie de forţă de muncă la lucrările de întreţinere.
La tomate, fasole şi mazăre există două tipuri de soiuri: cu creştere nedeterminată
(prezintă în vârf un mugure vegetativ, fructele apar şi cresc concomitent cu
creşterea tulpinii) şi cu
creştere determinată (în vârf, la un moment dat, mugurele vegetativ se transformă
în mugure florifer
şi se limitează creşterea în înălţime a plantei).
La ardei, la punctul de ramificare al tulpinii apare de obicei o floare, care la
culturile din
sere se recomandă să fie îndepărtată încă din faza de boboc, pentru ca
ramificaţiile să crească cât
mai uniforme, dând posibilitatea alegerii a 3 - 4 mai viguroase, pentru conducerea
plantelor.
La conopidă, spanac, salată, ceapă verde etc., zilele scurte şi temperaturile mai
scăzute
determină obţinerea părţilor comestibile de calitate şi cu perioadă mai lungă de
menţinere în acest
stadiu. Din contră, zilele lungi şi temperatura ridicată favorizează trecerea
rapidă a plantelor din
faza de creştere vegetativă în faza de fructificare, iar calitatea părților
comestibile scade simţitor.
Păstrarea usturoiului la temperaturi ridicate (peste 20°C) face ca plantele să nu
formeze bulb;
răsadul de ceapă plantat întârziat (la începutul îngroşării bulbului) nu mai
formează bulb; arpagicul
păstrat la 3 - 5°C sau la 15 - 18°C nu formează bulbi, doar tulpini florifere
(fuşti).
Test de autoevaluare
Având în vedere noţiunile parcurse în această unitate de învăţare,
răspundeţi la următoarele întrebări:
Întrebarea nr. 1
a) Care sunt speciile rezultate, plecând de la specia sălbatică, care au cel
mai mare grad de variabilitate?
b) Ce faze cuprinde perioada de sămânţă?
32c) Ce faze cuprinde perioada de creştere vegetativă?
d) Ce faze cuprinde perioada de reproducere?
e) Care sunt particularităţile creşterii şi dezvoltării la unele specii
legumicole şi ce importanţă au acestea pentru practică?
După parcurgerea acestei unităţi de învăţare trebuie să reţineţi:
 cum au evoluat speciile legumicole de-a lungul timpului şi ce factori au
influenţat evoluţia acestora;
 procesele specifice fiecărei perioade şi faze din viaţa unei plante şi ce
implicaţii tehnologice au acestea;
 care sunt particularităţile proceselor de creştere şi dezvoltare a
plantelor legumicole şi relaţia acestora cu tehnologia de cultură.
3.5. Comentarii şi răspunsuri la teste
Întrebarea nr. 1
a) Speciile legumicole care au rezultat în urma selecţiei pornind de la
specia sălbatică sunt varza albă, varza roşie, varza de Bruxelles, varza
creaţă, varza de cocean, conopida, broccoli, gulia, varza de frunze.
Aceste specii au apărut ca urmare a selecţiei îndelungate a formelor
valoroase, procesul fiind influenţat şi de factorii climatici.
Variabilitatea mai este pregnantă şi la alte specii legumicole, precum:
ridichile, castraveţii, dovlecii, tomatele, ardeiul, vinetele etc.
b) Perioada de sămânţă cuprinde 3 faze, şi anume: faza embrionară care
începe din momentul fecundării şi durează până la maturarea
seminţelor, în care organismele sunt foarte sensibile la schimbări şi
sunt dependente de planta mamă; faza de repaus durează din momentul
în care seminţele devin mature din punct de vedere fiziologic, până
când se declanşează procesul de germinare al acestora, procesele
metabolice sunt mult încetinite - cuprinde 2 subfaze: repaosul profund
şi repausul forţat; faza de germinare începe din momentul crăpării
tegumentului seminal şi ţine până la formarea primei frunze adevărate.
Plantele manifestă cerinţe diferite faţă de temperatură şi umiditate şi
germinează într-un anumit număr de zile în funcţie de specie.
33c) Perioada de creştere vegetativă cuprinde: faza de răsad care durează de
la apariţia primei frunze adevărate şi până la începerea depunerii
substanţelor de rezervă în organele specializate. Factorii de mediu
trebuie dirijaţi şi corelaţi în funcţie de cerinţele fiecărei specii. Hrana
absorbită este folosită în exclusivitate pentru creştere, raportul dintre
asimilaţie şi dezasimilaţie este aproape unitar; faza de acumulare
începe din momentul în care surplusul de hrană sintetizat de plante, se
depune în organe specializate, acestea cresc în dimensiuni, îşi modifică
forma şi sunt diferite în funcţie de specie; faza de repaus este specifică
plantelor bienale, trienale şi multianuale şi începe din momentul în care
metabolismul plantei se reduce foarte mult datorită factorilor de mediu
nefavorabili. Rădăcinile active mor, cu excepţia speciilor perene, la
care rămân active.
d) Perioada de fructificare cuprinde, de asemenea, 3 faze: faza de
îmbobocire care începe odată cu apariţia bobocilor florali şi se încheie
cu maturarea celulelor sexuale (a gameţilor). Compoziţia organelor
comestibile se schimbă, de aceea la plantele de la care se recoltează
părţile comestibile, recoltarea trebuie să se facă înaintea declanşării
acestei faze; faza de înflorire durează din momentul maturării
gameţilor până în momentul fecundării şi influenţează foarte mult
alegerea momentului optim pentru hibridare; faza de fructificare
începe din momentul fecundării şi durează până când seminţele devin
mature din punct de vedere fiziologic şi independente de planta mamă.
Face trecerea de la o generaţie la alta.
e) Particularităţi de creştere şi dezvoltare la tomate: formarea rădăcinilor
pe tulpină în contact cu solul umed, ceea ce permite plantarea mai
adâncă şi înmulţirea prin butaşi; existenţa soiurilor cu creştere
determinată, folosite pentru cultura destinată industrializării, la care
creşterea în înălţime este determinată genetic şi se manifestă prin
transformarea la un moment dat a mugurelui vegetativ în mugure
florifer, nu se copilesc, nu se palisează; soiuri cu creştere
nedeterminată, care au în vârf un mugure vegetativ care asigură
creşterea în înălţime, cât factorii de mediu sunt favorabili; se foloses
pentru cultura în sere, solarii, în câmp (cultură timpurie) şi obligatoriu
se palisează şi se copilesc; polenizarea este autogamă şi alogamă.
La castraveţi există hibrizi ginoici care formează numai flori femele
atât pe tulpina principală, cât şi pe lăstari, nu necesită ciupire în
vederea apariţiei lăstarilor de ordin superior, sunt foarte productivi. La
unele soiuri este specific fenomenul de partenocarpie (fructe fără
seminţe).
343.6. Lucrare de verificare
Lucrarea de verificare solicitată implică activităţi care necesită
cunoaşterea Unităţii de învăţare nr. 3.
Răspunsurile la întrebări vor fi transmise tutorelui pentru comentarii,
corectare şi evaluare.
Pe prima pagină a lucrării se vor scrie următoarele: titulatura acestui curs
(LEGUMICULTURĂ GENERALĂ), numărul lucrării de verificare, numele şi
prenumele studentei (studentului).
Fiecare răspuns va trebui să fie clar exprimat şi să nu depăşească o
jumătate de pagină. Punctajul aferent este menţionat pentru fiecare întrebare.
Întrebările la care trebuie să răspundeţi sunt următoarele:
1. Care sunt centrele de origine ale plantelor legumicole? – 1 p.
2. Cum explicaţi variabilitatea speciilor legumicole? – 1 p.
3. Caracterizaţi perioada de sămânţă. – 2 p.
4. Caracterizaţi perioada de creştere vegetativă. – 2 p.
5. Care sunt particularităţile creşterii şi dezvoltării la tomate şi castraveţi
şi cum influenţează tehnologia de cultură? – 3 p.
În ultima parte a lucrării, vă rog să comentaţi conţinutul testelor de
autoevaluare şi să scrieţi ce credeţi că ar trebui să cuprindă acestea pentru a fi
mai eficiente.
* Un punct se acordă din oficiu.
3.7. Bibliografie minimală
1. Ceauşescu I. şi colab., 1984 – Legumicultură generală şi specială. EDP,
Bucureşti.
2. Hoza Gheorghiţa, 2011 – Legumicultură generală. Editura Ceres, Bucureşti.
3. Indrea D., Apahidean A.Al., Maria Apahidean, Mănuţiu D., Rodica Sima, 2007 –
Cultura legumelor. Editura Ceres, Bucureşti.
35Unitatea de învăţare nr. 4
SUBSTANŢELE BIOACTIVE
ŞI CLASIFICAREA PLANTELOR LEGUMICOLE
Cuprins
4.1. Obiectivele unităţii de
învăţare...........................................................................
........ 36
4.2. Substanţe bioactive folosite în
legumicultură............................................................. 36
4.2.1. Substanţe
stimulatoare.......................................................................
..............37
4.2.2. Substanţe
retardante.........................................................................
............... 41
4.2.3. Substanţe
inhibitoare........................................................................
............... 41
4.2.4. Substanţe
adjuvante..........................................................................
............... 43
4.3. Clasificarea plantelor
legumicole.........................................................................
...... 44
4.4. Comentarii şi răspunsuri la
teste..............................................................................
... 51
4.5. Lucrare de
verificare.........................................................................
.......................... 52
4.6. Bibliografie
minimală...........................................................................
...................... 52
4.1. Obiectivele unităţii de învăţare
La sfârşitul acestei unităţi de învăţare studenţii vor fi capabili să:
 cunoască tipurile de substanțe bioactive și modul de acțiune asupra
plantelor legumicole;
 cunoască criteriile de clasificare a plantelor legumicole și caracterizarea
acestora.
4.2. Substanţe bioactive folosite în legumicultură
Substanţele bioactive în mare parte sunt sintetizate de plante, dar sunt obţinute
şi pe cale
chimică, prin sinteză, cu efecte foarte asemănătoare cu a celor naturale.
Aceste substanţe se
împart în următoarele
grupe:




substanţe stimulatoare;
substanţe retardante;
substanţe inhibitoare;
substanţe adjuvante.
364.2.1. Substanţe stimulatoare
În grupa substanţelor stimulatoare sunt cuprinse:
- auxinele;
- giberelinele;
- citochininele.
Auxinele sunt substanţe care controlează în principal procesul de creştere al
plantelor.
Acestea pot fi endogene (naturale), sintetizate de către plante şi acumulate în
organele tinere
(muguri, vârfuri de creştere). Pe baza auxinelor endogene au fost obţinute pe cale
chimică o serie de
compuşi asemănători ca structură şi ca mod de acţiune cu acestea.
Rolul auxinelor este de a stimula creşterea în înălţime prin diviziunea şi
elongaţia celulelor,
formarea rădăcinilor şi creşterea capacităţii de absorbţie a apei şi elementelor
minerale, creşterea
fructelor, îngroşarea membranelor celulare, stimularea înrădăcinării butaşilor la
speciile legumicole
care se înmulţesc prin butaşi (batat), germinarea seminţelor şi stimularea
fructificării.
Dintre auxinele sintetizate pe cale artificială, o folosire mai largă au
următoarele: ANA
(acidul naftil acetic), AIA (acidul indolil acetic), IBA (acidul indolil butiric),
BIB (beta indolil
butiric). Aceste substanţe se folosesc în doze foarte mici pentru a avea efectul de
stimulare a unor
procese, în concentraţii mari pot deveni toxice. Astfel, AIA şi BIB se folosesc în
concentraţie de 1 -
200 mg/l, ANA 1 - 10 mg/l etc.
Giberelinele sunt substanţe cu acţiune stimulatoare, descoperite şi izolate din
ciuperca
Gibberella fujikuroi. Giberelinele au fost depistate şi în seminţele şi rădăcinile
speciilor
leguminoase (mazăre, fasole), în tuberculii de cartof şi în seminţele imature de
pepene verde. Pe
cale artificială s-au creat până în prezent 23 de gibereline, cu acţiune
asemănătoare celor sintetizate
de către plante şi au fost notate cu GA de la 1 la 23, însă cele mai folosite sunt
GA 3 , GA 1 , GA 4 şi
GA 2 . Acţiunea giberelinelor se manifestă prin accelerarea unor procese
metabolice, contribuie la
sintetizarea şi acumularea de auxine endogene, stimulează germinarea seminţelor,
creşterea
aparatului foliar, inducerea partenocarpiei la vinete, schimbarea raportului între
florile femele şi
florile mascule la cucurbitacee, provoacă înflorirea unor specii bienale (morcov,
pătrunjel etc.) în
primul an de cultură, determină modificări ale metabolismului plantelor (scade
conţinutul în
amidon, azot total şi proteine şi creşte conţinutul în acizi nucleici), se
intensifică unele fenomene
fiziologice (fotosinteza, respiraţia) ca urmare a creşterilor active şi a sporirii
suprafeţei foliare etc.
În urma tratării plantelor cu gibereline, consumul de apă creşte, iar plantele
înregistrează o
sensibilitate mai mare la secetă.
Citochininele sunt substanţe stimulatoare sintetizate în rădăcini. Prima
citochinină a fost
identificată de Letham (1964) în seminţele imature de porumb şi a fost numită
zeatină. Rolul
citochininelor este de a preveni sau întârzia îmbătrânirea ţesuturilor, stimularea
creşterii prin
extensie şi diviziune celulară, formarea florilor şi a fructelor partenocarpice,
înlăturarea dominanţei
apicale, creşterea rezistenţei plantelor la stres (termic, hidric), la atacul
bolilor şi dăunătorilor. Pe
cale artificială s-au obţinut citochinine sintetice cu acţiune foarte asemănătoare
cu a kinetinei. Cele
mai folosite citochinine sunt: 1-2 difenilurea, 1-benzil-adenina, 8-a azochinetina
etc.
Acţiune stimulatoare manifestă şi vitaminele asupra germinării seminţelor,
absorbţiei apei
şi elementelor minerale, creşterii rezistenţei plantelor în condiţii de mediu mai
precare, creşterii
producţiei şi îmbunătăţirii calităţii fructelor etc. Dintre vitamine un rol mai
important joacă
vitaminele din complexul B, vitamina C şi PP.
37Produsele comerciale
Biostimulatori de înrădăcinare
Sprintene se foloseşte pentru stimularea creşterii sistemului radicular, prin
înrădăcinarea
mai profundă a plantelor şi creşterea capacităţii de ramificare, reducerea
stresului transplantării prin
formarea rapidă de rădăcini noi, adaptarea mai uşoară la noile condiţii de sol. Se
aplică radicular şi
foliar, în diferite fenofaze, şi anume: înainte de semănat prin umectarea
seminţelor timp de 2 - 4 ore,
în soluţie cu concentraţia de 0,1 %, după răsărire pentru a stimula creşterea
frunzelor şi a rădăcinilor
în primele faze 0,1 %, după plantare, odată cu prima udare, folosind 3 - 5 l/ha,
iar dacă aplicarea nu
s-a realizat în condiţii optime, tratamentul se repetă după o săptămână. Când
plantele sunt afectate
de stres termic, datorat fie de temperatura ridicată şi insolaţia puternică, fie de
temperatura scăzută
(îngheţ), se aplică 3 tratamente săptămânale, folosind 5 l/ha, stimulând refacerea
acestora.
Radifarm este un extract vegetal care conţine polizaharide, proteine şi
polipeptide,
îmbogăţit cu aminoacizi, vitamine şi kelaţi de fier şi zinc. Are rol stimulator în
formarea şi
ramificarea sistemului radicular prin stimularea sintezei hormonilor de la nivelul
rădăcinilor,
activează metabolismul plantelor prin acţiunea vitaminelor şi microelementelor,
plantele suportă
mai uşor stresul transplantării, stimulează fructificarea şi întregul proces de
dezvoltare al plantelor.
Se recomandă să se aplice două tratamente, cu condiţia ca soluţia să ajungă în zona
rădăcinilor.
Primul se aplică la repicat prin scufundarea rădăcinilor sau udarea ghivecelor cu o
soluţie de
Radifarm 0,3 %, iar al doilea prin distribuire odată cu apa de irigat (500 ml
Radifarm/1.000 mp) sau
prin udare locală, cu o soluţie în concentraţie de 0,25 %. Se foloseşte la tomate
şi ardei, dar nu
numai.
Radistim 1 este un stimulator de înrădăcinare folosit pentru butaşii erbacei
(tomate, batat,
tarhon etc.), a cărui reacţie se manifestă prin creşterea procentului de butaşi
înrădăcinaţi, numărul şi
lungimea rădăcinilor, protejarea de diferiţi agenţi patogeni, vigoare mai mare a
plantelor. Se
prezintă sub formă de pudră, în care se introduce baza butaşilor, dar pentru a avea
efect, punerea
butaşilor în substrat trebuie să se facă cu atenţie, pentru ca produsul să rămână
cât mai mult pe baza
acestuia. Poate fi şi sub formă lichidă.
Razormin este un biostimulator pentru înrădăcinare, conţine macro şi microelemente,
aminoacizi, polizaharide, care determină o creştere foarte bună a sistemului
radicular, cu implicaţii
pozitive asupra creşterii vegetative şi a fructificării plantelor. Induce absorbţia
la nivelul sistemului
radicular a nutrienţilor din sol, protejează plantele când se află în stare de
fitotoxicitate sau de stres,
are efect revitalizant şi prelungeşte perioada de vegetaţie, îmbunătăţeşte
calitatea fructelor prin
culoare şi conţinut în zaharuri, accelerează activitatea microbiologică a solului.
Se aplică pe
întreaga perioadă de vegetaţie, în concentraţie de 0,05 - 0,1 %.
Biorootz stimulează creşterea numărului de microorganisme din sol, care provoacă o
absorbţie mult mai mare a nutrienţilor, precum şi protecţia la diverse boli ale
solului; mai mult,
acest produs are efect inhibitor asupra mucegaiurilor, limitează răspândirea
virusurilor, reduce
poluarea, creşte capacitatea de absorbţie a îngrăşămintelor de către plantă cu
circa 40 %.
Revital se foloseşte pentru stimularea formării rădăcinilor şi trecerea mai uşoară
peste
stresul transplantării la tomate, ardei şi vinete. Produsul se foloseşte în
concentraţie de 0,05 - 0,1 %.
Aplicat în faze timpurii de vegetație, accelerează şi intensifică creşterea
rădăcinilor
secundare care au rol important în procesul de absorbţie a apei şi substanţelor
minerale, asigură o
mai bună prindere a răsadurilor după repicat şi plantare, plantele sunt mai
viguroase, fructificarea
este mai abundentă, creşte rezistenţa la boli şi la diferiţi factori de stres, în
special temperatura
excesivă.
38Tecamin raiz. Prin conţinutul în macroelemente, microelemente, aminoacizi şi alge
marine îmbunătăţeşte procesele de germinare şi răsărire a plantelor, stimulează
ramificarea
rădăcinilor şi formarea perişorilor absorbanţi. Aplicarea produsului se face prin
instalaţia de
picurare sau prin procedeul de fertirigare, la 5 - 7 zile după ce plantele au
răsărit, cu o soluţie în
concentraţie de 0,1 - 0,2 %. La plantare răsadurile se imersează într-o soluţie de
1 %, asigurând o
prindere foarte bună şi o uniformitate ridicată a culturii din primele faze.
Biostimulatori de creştere
Cropmax este stimulator de creştere natural, care poate fi folosit la culturile din
câmp, sere
şi solarii. Conţine macroelemente, microelemente, vitamine, aminoacizi, hormoni şi
enzime
vegetale. Se foloseşte atunci când în sol este un conţinut scăzut de NPK şi
microelemente, pH
ridicat, temperatură scăzută şi activitate slabă a rădăcinilor, dezechilibru în sol
între K, Ca şi Mg sau
când solul este slab aerat. Se aplică prin pulverizare fină, recomandat pe ambele
feţe ale frunzei
pentru o mai bună absorbţie şi în perioada din zi când activitatea fotosintetică
este optimă. Se
recomandă 1 - 3 tratamente în timpul perioadei de vegetaţie, folosind 500 - 1.000
ml/ha la fiecare
tratament. Se poate aplica şi săptămânal, în concentraţie de 0,2 %, obţinând
importante sporuri de
producţie. Este compatibil cu majoritatea pesticidelor, mai puţin cu cele pe bază
de cupru.
Atonik este un stimulator de creştere care se absoarbe şi se translocă foarte rapid
în plantă,
intensifică translocarea sevei brute şi asimilatelor, creşte conţinutul în
clorofilă, determină creşterea
lăstarilor, înflorirea şi fructificarea, stimulează înrădăcinarea butaşilor,
germinarea seminţelor,
multiplicarea microorganismelor din sol şi contribuie la descompunerea materiei
organice,
ameliorând fertilitatea solului. Se foloseşte pentru umectarea seminţelor, tratarea
bazei butaşilor
pentru o mai bună înrădăcinare şi pentru stimularea înfloririi şi fructificării.
Plantele tratate cu
Atonik dau producţii mai mari, mai timpurii şi de calitate foarte bună. Se aplică
de 2 - 4 ori în cursul
unei perioade de vegetaţie, în funcţie de specie şi sistemul de cultură, în
Mai concret, se aplică la ardeiul cultivat în solarii, pentru stimularea creşterii
sistemului
radicular, în concentraţie de 0,025 %; la cartof, pentru stimularea înfloririi, 0,5
l/ha; la castraveţi,
pentru obţinerea de producţii timpurii în concentraţie de 0,05 - 0,025 % (2,5
l/ha); la tomatele
timpurii şi cultivate în solar, pentru creşterea producţiei, în concentraţie de
0,025 % (2,5 l/ha).
Maxicrop start conţine chelaţi Mg, Mn, Fe, Zn şi substanţe active selecţionate din
extracte
vegetale şi alge marine (100 % Ascophyllum nodosum destinat stimulării creşterii
vegetative). Se
aplică numai foliar, de 2 ori, de la repicat până la înflorire, în concentraţie de
0,15 - 0,2 %.
Kendal este un biostimulator care are în compoziţie extracte naturale,
oligozaharide, săruri
de potasiu şi glutation cu acţiune de autoapărare a plantelor şi de nutriţie, care
creşte mecanismul de
apărare natural al plantelor, stimulează procesele de creştere şi dezvoltare a
plantelor, previne şi
reduce atacul de ciuperci şi bacterii patogene. Se aplică foliar, la interval de 10
- 15 zile, în cantitate
de 1,5 - 2 l/ha. Se poate aplica şi local în concentraţie de 0,3 - 0,4 %, circa 100
- 200 ml soluţie/plantă.
Biostimulatori de legare a fructelor
Auxigib este un fitoregulator sub formă de pulbere umectabilă, folosit pentru a
stimula
înflorirea rapidă şi concentrată, fructificarea şi partenocarpia. De asemenea,
contribuie la revenirea
plantelor afectate de temperaturi scăzute şi diminuarea pagubelor. Se aplică la
tomate, în mod
repetat, în timpul înfloririi, prin pulverizări asupra plantelor, cu soluţii în
Pentru revenirea plantelor afectate de ger (cele care iernează sub formă de
rozetă), imediat după
îngheţ, dar înaintea începerii dezgheţului, se aplică stropiri cu soluţii în
concentraţie de 0,75 - 0,85 %,
cu efecte benefice asupra acestora.
3936 c tipo b este fitoregulator de legare, favorizează fecundarea florilor, reduce
căderea
prematură a florilor şi nu determină deformarea fructelor. Se foloseşte la tomate
şi vinete, prin
pulverizare foarte fină pe flori, când sunt deschise, în cantitate de 1 - 5 ml/l,
la pepenele galben 1 -
2 ml/l, prin pulverizarea întregii plante, dar fără exces. Când se aplică la
hibrizi, cantitatea de
produs trebuie să fie mai mică.
Adrop este un fitoregulator cu acţiune de legare a fructelor, stimulator şi
cicatrizant.
Produsul, bazat pe trei substanțe active de natură auxinică, poate fi folosit la o
gamă largă de culturi,
determinând precocitatea, creşterea şi uniformizarea fructificării, creşterea
fructelor, reduce
avortarea florilor, stimulează creşterea vegetativă şi productivă a culturii, are o
acţiune cicatrizantă
asupra ţesuturilor vegetale lezate. Reduce pagubele datorate gerului, prin
aplicarea unui tratament
preventiv cu 24 ore înaintea apariţiei gerului şi se repetă dacă fenomenul persistă
sau imediat după
apariţia acestuia. Se aplică sub formă de soluţii în concentraţie de 0,5 - 0,7 %.
Stimolante 66 F este un stimulator vegetal care conţine activatori ai funcţiilor
metabolice,
provocând dezvoltarea plantelor şi îmbunătăţirea caracteristicilor calitative şi
cantitative ale
producţiei. Ajută la parcurgerea mai uşoară a fenofazelor de creştere şi
fructificare, când plantele
sunt în stare de stres şi stagnare fiziologică datorită îngheţului, aplicării
tratamentelor fitosanitare în
exces sau când acestea nu s-au aplicat.
Betagib LG este un fitoregulator folosit pentru legarea florilor la vinete. Se
foloseşte sub
formă de soluţie, care se aplică când floarea este deschisă, prin pulverizare fină,
în concentraţie de
0,9 -1 %.
Maxicrop SET conţine chelaţi de B şi Zn şi substanţe active selecţionate din
extracte
vegetale și alge marine (100 % Ascophyllum nodosum), este destinat stimulării
proceselor de
înflorire şi legare a fructelor. Se aplică numai foliar, în perioada cuprinsă între
înflorire şi legarea
fructelor, de 1 - 2 ori, în concentraţie de 0,15 - 0,2 %.
Biostimulatori pentru maturarea fructelor
Maxicrop quality conţine chelaţi de B, Mn, Fe, Ca şi substanţe active selecţionate
din
extracte vegetale și alge marine (100 % Ascophyllum nodosum), destinat stimulării
maturării
fructelor şi obţinerii de fructe calitativ superioare, în ceea ce priveşte gustul,
culoarea şi consistenţa.
Se aplică foliar, la interval de 7 - 10 zile, începând de la formarea fructelor,
până la recoltare, în
Brixer este folosit pentru a concentra şi a uniformiza maturarea, reducând numărul
de
treceri de recoltare. Are şi efect uşor desicant asupra resturilor vegetale, când
recoltarea se face o
singură dată. Conţine substanţe naturale (metianină, polizaharide, extrase de alge
şi yucca) care
accelerează procesele de maturare a fructelor, favorizează anticiparea coloraţiei
şi determină
acumularea zaharurilor. Aplicat pe cale foliară, prin pulverizare, îmbunătăţeşte
substanţial nivelul
calitativ al fructelor. Se recomandă două tratamente, primul cu 30 de zile înainte
de recoltare, iar al
doilea după 15 zile de la primul, în concentraţie de 0,2 - 0,3 %.
Gerephon SL este un fitoregulator de sinteză pe bază de etefon, care după ce este
absorbit
în plantă, eliberează etilena, gaz produs în mod normal de către plantă,
determinând stimularea
maturării fructelor. Se foloseşte în special la tomate, atât pentru
industrializare, cât şi pentru consumul
în stare proaspătă, cu scopul anticipării şi uniformizării maturării fructelor.
Administrarea
produsului se face atunci când 40 - 60 % din fructe s-au maturat, iar cele din
ultima inflorescenţă au
atins mărimea caracteristică cultivarului şi au culoarea verde, în cantitate de 3 -
4 l/ha. Când se
doreşte concentrarea maturării, condiţionată de eliberarea terenului, pentru
recoltarea fructelor mari
şi mici, se aplică pulverizarea întregii plante, cu o soluţie în concentraţie de
0,2 - 0,25 %.
404.2.2. Substanţe retardante
Spre deosebire de substanţele stimulatoare care sunt naturale şi artificiale,
substanţele
retardante sunt în exclusivitate substanţe chimice. Acestea se produc la scară
industrială, rolul lor
asupra plantelor fiind de încetinire o anumită perioadă de timp a creşterii în
înălţime, fără să
afecteze frunzele, florile sau fructele. Substanţele retardante acţionează asupra
procesului de
elongaţie a celulelor, stopând alungirea tulpinilor. Se aplică întotdeauna înaintea
alungirii plantelor,
deci preventiv, şi în general la speciile cu un ritm accelerat de creştere,
predispuse alungirii. În
practica legumicolă se aplică pe scară largă la tomate.
Produse comerciale
Cycogan se aplică la tomate în faza de răsad, odată sau de două ori, primul
tratament
efectându-se când plantele au 3 - 4 frunze bine formate. Concentraţia soluţiei de
Cycogan este de
0,1 - 0,15 %, se distribuie pe plante prin pulverizare foarte fină, cu aparate de
stropit portabile. În
urma tratării tomatelor cu Cycogan, plantele sunt mai viguroase, cu tulpina mai
scurtă şi mai groasă,
internodurile mai scurte şi cu o rezistenţă mult mai bună la temperaturi scăzute.
Ridichiile de lună
în faza de 2 frunze, tratate cu produse similare (Cycocel) în concentraţie de 1.000
- 2.000 ppm au
realizat un spor de producţie de 130 %, la varză s-au obţinut căpăţâni mai îndesate
şi o producţie
mai mare, la pepenele galben o creştere a numărului de flori femele, sporuri de
producţie la
rădăcinoase (Stan N., 1999) etc.
Cycogan 40 LC în concentraţie de 0,05 - 0,1 % folosit la tomate şi ardei asigură
stimularea
creşterii răsadurilor şi maturarea fructelor.
Stabilan este un produs omologat pentru grâu, rapiţă şi tomate, care se aplică prin
pulverizare fină şi acţionează asupra taliei plantelor. Absorbţia produsului este
destul de rapidă,
circa 2 - 4 ore şi este condiţionată de umiditatea ridicată. La tomate, se aplică
la răsaduri, având ca
efect obţinerea de răsaduri de calitate mai bună, cu tulpini şi internoduri mai
scurte, cu capacitate
mai bună de fructificare şi fructe de calitate superioară. Se aplică preventiv, în
concentraţie de 0,1 %,
câte 1 litru/mp.
Alar se foloseşte la culturile de tomate, ardei şi vinete în perioada de răsad, în
concentraţie
de 0,3 % când acestea au 3 - 4 frunze, prin pulverizare fină pe frunze, prevenind
alungirea acestora.
Se poate aplica şi în cultură, în concentraţie de 0,25 %, efectuând 3 tratamente,
primul la 2 - 3
săptămâni de la plantare şi mai ales în perioada intrării fructelor în pârgă, la
interval de 2 - 3
săptămâni, pentru stimularea înfloririi, creşterii procentului de legare a
fructelor şi maturarea
concentrată a acestora.
4.2.3. Substanţele inhibitoare
Sunt substanţele chimice sau naturale care stopează acţiunea substanţelor
stimulatoare,
mergând până la oprirea creşterii plantelor şi intrarea acestora în repaus.
o
o
o
o
Substanţe inhibitoare naturale
Abscisin
Cumarina
Acidul cinamicnd iar să nu
poată oferi pe piaţa agrară decât forţa braţelor denger and ends the
reaction. (Diagrams after Lodish et al., 2000.) Many cell surface receptors act
indirectly. When
they bind to a ligand they induce a series of in-
tracellular activation steps. This reaction sys-
tem consists of a receptor protein, a protein (G
protein) bound to a guanosine residue, and an
enzyme to be activated. Ligand binding alters
the receptor protein and activates the G protein
(2). This moves to the effector, e.g., an enzyme
complex (3), and activates it (4). In this way, a
second messenger is formed that triggers
further reactions in the cell, e.g., cyclic ade-
nosine monophosphate (cAMP) by means of the
enzyme adenylate cyclase (see p. 268).
References
Lodish, H. et al.: Molecular Cell Biology. 4 th ed.
Scientific American Books, New York, 2000.
Watson, J.D., et al.: Recombinant DNA. 2 nd ed.
B. Hormones with immediate effects
on cells
Important examples of hormones that function
as ligands are amino acid derivates, arachidonic
acid derivatives, and many peptide hormones.
All rights reserved. Usage subject to terms and conditions of license.Types of Cell
Surface Receptors
267268
G Protein-coupled Receptors
The indirect transmission of signals into the cell
is mediated by transmembrane proteins, which
traverse the cell membrane. A first messenger,
e.g., a hormone like epinephrine, triggers an in-
tracellular reaction by binding to a specific re-
ceptor. This leads to activation of a second mes-
senger, which in turn initiates a series of reac-
tions that result in a change of cell function.
Many of the genes for the different proteins in-
volved in the indirect transmission of signals
are known.
A. Stimulatory G protein (G s ) and
hormone–receptor complex
There are many endogenous messengers (hor-
mones) with their own specific receptors. First
the hormone binds to the receptor (formation
of a hormone–receptor complex). The intra-
cellular transmission of signals is mainly car-
ried out by special guanine-nucleotide-binding
proteins, or G proteins. By binding to guanosine
triphosphate (GTP, a nucleotide composed of
guanine, a sugar, and three phosphate groups),
the G protein becomes activated and initiates
further reactions. G proteins consist of three
subunits: α , β , and γ . The α subunit (stimulatory
G protein, G s ) binds to the effector protein. Im-
mediately thereafter, G α is inactivated (GTPase)
by hydrolysis of GTP to GDP (guanosine di-
phosphate). This transforms the G protein back
into an inactive form (G i ).
Several human diseases due to defective G pro-
tein or a defective G protein receptor are known.
(Clapham, 1993.)
B. Four hormone classes
Four principal classes of hormones can be
differentiated: (1) amino acid derivatives such
as epinephrine and epinephrine derivatives; (2)
polypeptides such as glucagon; (3) steroids
such as cortisol and its derivatives; and (4) fatty
acid derivatives such as the prostaglandins.
sponsible for many physiological reactions. It
becomes inactivated when converted into ade-
nosine monophosphate (AMP) by phos-
phodiesterase. cGMP (cyclic guanosine mono-
phosphate) functions in the same manner as
cAMP to initiate an intracellular reaction.
D. G protein cycle to activate adenylate
cyclase
When a hormone binds to its specific receptor, a
structural change occurs (1). This activates the
α subunit of the G protein, which separates
from the β and γ subunits (2). The stimulatory G
protein (G s - α ) binds to the effector protein, usu-
ally adenylate cyclase, and activates it (3). cAMP
is then formed from ATP, while GTP is hydro-
lyzed to GDP at the G- α subunit. This inactivates
the effector protein and the formation of cAMP
is terminated. Thus, the signal is of very short
duration, and the initial conditions are rapidly
restored. Several toxins exert their activity by
interrupting this cycle. For example, cholera
toxin inhibits inactivation of the G s - α protein so
that adenylate cyclase remains activated and
large amounts of sodium and water are lost
through the intestinal mucous membranes.
(Figures adapted from Watson et al., 1992).
References
Bourne, H.R., Sanders, D.A., McCormick, F.: The
GTPase superfamily: conserved structure
and molecular mechanism. Nature 349 :11 –
127, 1991.
Clapham, D.E.: Mutations in G protein-linked
receptors: novel insights on disease. Cell
75 :1237 – 1239, 1993.
Linder, M.E., Gilman, A.G.: G-Proteins, Sci. Am.
36 – 43, 1992.
Watson, J.D., et al.: Recombinant DNA. 2 nd ed.
W.H. Freeman, Scientific American Books,
New York, 1992.
C. Formation and hydrolysis of cAMP
The key reaction is the formation of cyclic ade-
nosine monophosphate (cAMP) from adenosine
triphosphate (ATP) by means of adenylate cy-
clase. Intracellular cyclic AMP transmits the ac-
tivation initiated by the hormone–receptor
complex without a molecule having passed
through the plasma membrane. cAMP is re-
G Protein-coupled Receptors
Hormone
H
Receptor
Lipid bilayer
H
Receptor
Receptor
"
G protein
inactive
"
G protein active (G s )
GTP
Binding to G protein
!
#
!
#
Hormone-receptor complex
H
Receptor
"
!
Inactivation
of G ! (GTPase)
Effect
on
effector
protein
!
#
Activation of G !
GDP
A. Stimulatory G protein (Gs) and hormone-receptor complex
1 Norepinephrine
Phosphodiesterase
Adenylate cyclase
(amino acid derivative)
Adenine
2 Glucagon
Adenine
Adenine
(polypeptide)
P
3 Cortisol
(steroid)
4 Prostaglandin A 2
P
P
Ribose
Adenosine
triphosphate (ATP)
(fatty acid derivative)
B. Four hormone types
#
G protein
Cyclic adenosine
monophosphate (cAMP)
Ribose
Adenosine
monophosphate
(AMP)
H
Receptor
"
P
Ribose
C. Formation and hydrolysis of cAMP
Effector protein
e.g., adenylate cyclase
H
P
Receptor
"
!
#
!
G s
GTP
1. G protein binds to hormone-receptor
1. complex
2. G protein activated
H
Receptor
Receptor
"
#
"
#
!
GDP
!
ATP
cAMP
G s inactivated by GTPase
4. G protein inactivated
3. Effector protein activated
D. G protein cycle to activate adenylate cyclase
Physio-
logical
effect270
Functional signals between cells are received by
transmembrane proteins as signal transmitters.
During evolution, relatively simple precursor
genes for such proteins gave rise to multiple
structurally and functionally related genes.
Their corresponding proteins serve to transmit
ions (sodium, potassium, calcium, chloride, and
others), as neurotransmitters, and for percep-
tion of light and odors, etc. Cloning of these
genes has yielded insight into the variety of
functions of transmembrane signal transmit-
ters. Their general structure can be traced back
to an evolutionarily conserved ancestral
molecule. follow. An especially common structural motif
is a transmembrane protein containing seven α
helices within the plasma membrane. The
amino end is extracellular; the carboxy end is
intracellular. Different oligosaccharide side
chains are usually bound to the extracellular
domains. The intracellular domains have bind-
ing sites for other molecules involved in signal
transmission. The seven-helix motif is the
characteristic structure of G protein-binding re-
ceptors (p. 268). As the G proteins themselves,
these receptors and their genes form a large
family with a long evolutionary history. In
yeast, they serve to discern the pheromones of
the mating types (p. 186); in higher organisms
they are the basis for transmitting signals of vi-
sion, smell, and taste (p. 278 – 286). (Figure re-
drawn from Stryer, 1995).
A. Transmembrane structure of
voltage-gated ion channels C. A receptor with two trans-
membrane protein chains, α and β
The direct flow of ions across the cell membrane
is regulated by ion channels. The transmem-
brane proteins, composed of several domains,
are arranged so that they form pores that can be
opened and closed. The simplest model is the
potassium channel (1). This membrane-bound
polypeptide contains six transmembrane
domains. The amino and the carboxy ends of
the protein lie within the cell. Changes in cell
membrane potential or voltage cause the chan-
nel to open (or close) in order to initiate (or ter-
minate) a brief flow of ions. Domain 4, which is
composed of polar amino acids, is crucial for the
flow of ions. Sodium and calcium ion channels
consist of four subunits (2) of similar structure,
each resembling a potassium channel. The simi-
larity is due to the common evolutionary origin
of their genes. The four subunits of the sodium
channel (3) are positioned to form a very nar-
row porelike passage, much narrower than a
potassium channel, through the plasma mem-
brane. Ion transport is brought about by mem-
brane depolarization (3 and 4). (Figure after
Watson et al., 1992.) The receptor for γ -aminobutyric acid (GABA)
utilizes two transmembrane protein subunits, α
and β . Both the amino and the carboxy ends are
extracellular. The two chains are coded for by
different genes. Several oligosaccharide side
chains are present on the extracellular side. The
β chain contains a phosphorylation site for
cAMP-dependent protein kinase.
(Figures adapted from Watson et al., 1992).
Transmembrane Signal
Transmitters
References
Sabatini, D.D., Adesnik, M.B.: The biogenesis of
membranes and organelles. pp. 459 – 553,
In: Scriver, C.R., et al., eds., The Metabolic
and Molecular Bases of Inherited Disease.
7 th ed. McGraw-Hill, New York, 1995.
Stryer, L.: Biochemistry, 4 th ed. Freeman Publ.,
San Francisco, 1995.
Watson, J.D. et al.: Recombinant DNA, 2 nd ed.,
B. Seven-helix structure of
transmembrane signal transmitters
Indirect transmission of signals is more
frequent than the direct transport of ions or li-
gand-gated impulse transmission. Here, the
transmembrane protein is involved only in the
first step of signal transmission. Further steps
All rights reserved. Usage subject to terms and conditions of license.Transmembrane
Signal Transmitters
271272
Receptors of Neurotransmitters
Impulses are relayed between nerve cells or be-
tween nerve and muscle cells by various trans-
mitter molecules (neurotransmitters). Their ef-
fects are further relayed by receptors in the cell
membrane. Receptors can be differentiated ac-
cording to their structure, which in turn deter-
mines their specificity.
A. Acetylcholine as a neurotransmitter
Cholinergic synapses convey the nerve impulse
from one nerve cell to another or from a nerve
cell to a muscle cell (motor endplate). Acetyl-
choline leads to postsynaptic depolarization
through the release of potassium ions (K + ) and
the uptake of sodium ions (Na + ). This process is
regulated by an acetylcholine receptor.
B. Acetylcholine receptors
The acetylcholine receptors are of two geneti-
cally and functionally different types. Phar-
macologically they can be differentiated ac-
cording to the effects of nicotine and muscarine.
The nicotine-sensitive acetylcholine receptor is
an ion channel for potassium and sodium. It
consists of five subunits: two α , one β , one γ ,
and one δ (1). Acetylcholine binds as a ligand to
the two α subunits. Each subunit consists of
four transmembrane domains (2). Each subunit
is encoded by its own gene (3). These genes
have similar structures and nucleotide base
sequences. The ligand-gated ion channel is an
example of direct transport without an inter-
mediate carrier. A mutation in the second trans-
membrane region has been shown to change
the ion selectivity from cations to anions (Galzi,
1992.)
The muscarine-sensitive type of acetylcholine
receptor is a protein that contians seven trans-
membrane subunits (4). Since each exists in the
form of an α helix, it is referred to as a seven-
helix transmembrane protein (p. 270). The
amino end (NH 2 ) lies extracellularly; the car-
boxy end (COOH), intracellularly. The trans-
membrane domains are connected by intra-
cellular and extracellular polypeptide loops (4).
Different domains of the whole protein are dis-
tinguished (5) according to location and the
relative proportion of hydrophilic and hydro-
phobic amino acids. The amino end and the car-
boxy end each form a domain just like the intra-
cellular (a–c), and extracellular portions (d–f).
The transmembrane domains located within
the plasma membrane (1 – 7) consist for the
most part of hydrophobic amino acids. The
structure of the gene product corresponds to
the general structure of the gene (6). The differ-
ent domains are coded for by individual exons.
The DNA nucleotide sequences within function-
ally similar domains are similar.
The seven-helix transmembrane motif occurs
in many receptors. The general structures of the
genes and of the gene products are very similar,
but they differ in their specificity of binding to
other functionally relevant molecules (G pro-
teins). They play a role not only as neu-
rotransmitters but also in the transmission of
light, odors, and taste. (Figures based on Wat-
son et al., 1992.)
References
Galzi, G.L.: Mutations in the channel domain of
a neuronal nicotinic receptor convert ion
selectivity from cationic to anionic. Nature
359 :500 – 505, 1992.
Watson, J.D., Gilman, M., Witkowski, J., Zoller,
M.: Recombinant DNA. 2 nd ed. W.H.
Freeman, Scientific American Books, New
York, 1992.
Receptors of Neurotransmitters
Presynaptic
Acetylcholine
+
+
K+
+ + -
- ++ +
+ + +
- -
-
- - - - -
K + High
Na + Low
Na +
Polarized
Postsynaptic
Depolarized
Cholinergic synapse (nerve/nerve or nerve/muscle)
A. Acetylcholine as neurotransmitter
Two types of acetylcholine receptors
Seven-helix transmembrane protein
bound to G proteins
(muscarine-sensitive)
Cation-specific channel
in muscle of vertebrates
(nicotine-sensitive)
1. Acetylcholine
1. binds to
1. !-subunits
1. (ligand
1. binding)
Cations
(K +, Na + )
$
"
!
extracellular
d
NH2
3
4.
1
#
2
e
5
4
f
6
7
intracellulär
a
5.
2. Each subunit
2. has four
2. transmembrane
2. domains
A
COOH
b
c
1 a 2 d 3 b 4 e 5 c 6 f 7
A
B
a - c
d - f
1 - 7
B
Amino end
Carboxy end
intracellular domains
extracellular domains
transmembrane domains (hydrophobic)
6. Gene structure (diagram)
3. One gene for each subunit:
2 for
1 for
1 for
1 for
!-subunits
"-subunit
#-subunit
$-subunit
Exons
5'
A
Introns
1 a 2 d 3 b 4 e 5 c 6 f 7
The different domains are encoded
by individual exons
B. Acetylcholine receptor
3'
B274
More than 20 different disorders due to defec-
tive ion channel proteins resulting from gene
mutations are known. Such disorders include
cystic fibrosis (see p. 276), the long-QT syn-
drome, a special type of deafness, hereditary
hypertension (Liddle syndrome), familial per-
sistant hyperinsulinemic hypoglycemia of in-
fancy, some hereditary muscle diseases, and
malignant hyperthermia (see p. 372), among
other disorders.
A. Long-QT syndrome, a genetic
cardiac arrhythmia
Congenital long-QT syndrome is characterized
by a prolonged QT interval in the electrocardio-
gram (more than 460 ms, corrected for heart
rate), sudden attacks of missed heart beats
(syncopes) or series of rapid heart beats (tor-
sades de pointes), and an increased risk for sud-
den death from ventricular fibrillation in
children and young adults.
B. Different molecular types of
long-QT syndrome
Prolongation of the QT interval in the electro-
cardiogram results from an increase in the du-
ration of the cardiac action potential (1). The
normal potential lasts about 300 ms (phases 1
and 2). The resting membrane potential (phase
3) is reached by progressive inactivation of cal-
cium currents and increasing depletion of
potassium currents, which repolarize the cell.
In phase 0 the cell is quickly depolarized by acti-
vated sodium currents following an excitatory
stimulus.
LQT1 accounts for about half of the patients
with long-QT syndrome. The gene for LQT2 en-
codes a 1195-amino-acid transmembrane pro-
tein responsible for the other major potassium
channel that participates in phase 3 repolariza-
tion (HERG stands for (human-ether-r-go-go-
related gene, a Drosophila homologue). LQT3, a
sodium channel protein, consists of four sub-
units, each containing six transmembrane
domains and a number of phosphate-binding
sites. Homozygosity for LQT1 (KVLQT1 gene
Genomic Imprinting
dies very
early
2
Androgenetic
1
normal
develop-
ment
Extra-
embryonic
tissues
3
Diploid zygote
Fetus absent
or stunted
Preimplantation
failure in most
Normal
Fetus normal
Preimplantation
dies
later
4
Fetus normal
until 40 somite
stage
Preimplantation normal,
extra-embryonic
tissues underdeveloped
Gynogenetic
A. The importance of two different parental genomes
Two paternal genomes
1. Hydatidiform mole
Two maternal genomes
2. Hydatidiform mole
3. Ovarian teratoma
4. Triploidy 69, XXX
B. Human embryonic development depends on presence of a maternal and a paternal
genome
1.
P
Somatic cells
XX and XY
Male
P
2.
P Paternal
M
Inactive Active
Active Inactive
P
M Maternal
Female
M
M
Imprint
erased
Primordial
germ cells
P
M
3.
Imprint
reset
Gametes
P
M
4.
Zygote
C. Genomic imprinting is established in early embryonic development
Imprint
established228
Fundamentals
X-Chromosome Inactivation
During early embryonic development of mam-
malian females, one of the two X chromosomes
becomes inactivated. The inactivation is in-
duced by a gene (XIST, X-inactivation-specific
transcript) on the proximal long arm of the X
chromosome, transcribed from the inactive X.
X inactivation is a mechanism to balance X-
chromosomal gene expression between female
and male cells.
A. X chromatin
In 1949, Barr und Bertram observed a stainable
appendage in the nucleus of nerve cells of
female (1 and 3) but not of male cats (2). The
authors named this structure as “sex chro-
matin”. Similar structures were found drum-
sticks in peripheral blood leukocytes (4) and
small peripheral bodies in the nuclei of fibro-
blasts and oral mucosa cells (5) in humans. Each
of these structures represents one of the two X
chromosomes and are referred to as X chro-
matin. (Figures 1 – 3 from Barr and Bertram,
1949).
B. Scheme of X inactivation
Random inactivation of most of the genes of one
of the two X chromosomes in female cells
occurs early in embryogenesis, at about day 21
in humans. In a given cell, it involves the X chro-
mosome of either maternal or paternal origin.
The inactivation pattern is normally irreversible
and stably transmitted to all daughter cells. The
expected distribution is usually 1 : 1. In rare in-
stances this may be skewed toward a preferen-
tial type of inactivation. In extreme cases this
may result in clinical manifestation in a heter-
ozygous female if the majority of cells contain
the mutation on the active X chromosome.
C. Mosaic pattern of expression
Female somatic tissues show a mosaiclike dis-
tribution of cells expressing just one of the two
alleles. In mice, X-chromosomal coat-color mu-
tants show a mosaic of light- and dark-colored
coat patches (1, after Thompson, 1965). In
humans, a similar distribution of normal and
absent sweat pores is seen in female heterozy-
gotes for hypohidrotic ectodermal dysplasia.
The sweat pores of hemizygotes are absent
(hypohidrosis). Fingerprints of female hetero-
zygotes show areas with normal sweat pores
(black points) and areas with absent sweat pores
(2a and 2b, from Passarge and Fries, 1973). In
cell cultures from female heterozygotes for
X-chromosomal HGPRT deficiency (hypo-
xanthine–guaninephosphoribosyltransferase)
the colonies are either HGPRT Ϫ or HGPRT + (3).
(From Migeon, 1971, with kind permission of the
author and publisher).
D. Exceptions from X-inactivation
Transcriptional silencing of X-chromosomal
genes in mammalian female cells is not
complete. Some genes escape inactivation and
are expressed from both the active and the inac-
tive X chromosome. The majority of these genes
have a homologue on the Y chromosome,
reflecting a common evolutionary origin. Panel
D shows genes that are expressed on the inac-
tive human X chromosome. Most are located at
the ends of the X chromosomes. (Figure
adapted from Brown et al., 1997).
E. X-inactivation profile
An analysis of 224 X-linked transcripts showed
that 34 escape inactivation (Carrel et al., 1999).
Of these, 31 map to the short arm of the X chro-
mosome. The expressed genes are open circles,
the inactivated genes filled circles. Several
genes in the pseudoautosomal regi
observations are correct, the future of life and the universe will be far bleaker.
In the last four years astronomers have reported evidence that the expansion of the
universe is not just continuing but is speeding up, under the influence of a
mysterious
"dark energy," an antigravity that seems to be embedded in space itself. If that is
true and
the universe goes on accelerating, astronomers say, rather than coasting gently
into the
night, distant galaxies will eventually be moving apart so quickly that they cannot
communicate with one another. In effect, it would be like living in the middle of a
black
hole that kept getting emptier and colder.
In such a universe, some physicists say, the usual methods of formulating physics
may
not all apply. Instead of new worlds coming into view, old ones would constantly be
disappearing over the horizon, lost from view forever.
Cosmological knowledge would be fragmented, with different observers doomed to
seeing different pieces of the puzzle and no single observer able to know the fate
of the
whole universe or arrive at a theory of physics that was more than approximate.
37"There would be a lot of things about the universe that we simply couldn't
predict," said
Dr. Thomas Banks, a physicist at the University of California at Santa Cruz.
And perhaps most important, starved finally of the energy even to complete a
thought or
a computation, the domain of life and intelligence would not expand, but constrict
and
eventually vanish like a dwindling echo into the silence of eternity. "I find the
fate of a
universe that is accelerating forever not very appealing," said Dr. Edward Witten,
a
theorist at the Institute for Advanced Study.
That is an understatement, in the view of Dr. Lawrence M. Krauss, an astrophysicist
at
Case Western Reserve University in Cleveland, who along with his colleague Dr.
Glenn
D. Starkman has recently tried to limn the possibilities of the far future. An
accelerating
universe "would be the worst possible universe, both for the quality and quantity
of life,"
Dr. Krauss said, adding: "All our knowledge, civilization and culture are destined
to be
forgotten. There's no long-term future."
Einstein's Last Laugh
Until about four years ago, an overwhelming preponderance of astronomers subscribed
to
the view that the cosmic expansion was probably slowing down because of the
collective
gravity of the galaxies and everything else in the universe, the way a handful of
stones
tossed in the air gradually slow their ascent. The only question was whether the
universe
had enough gravitational oomph to stop expanding and bring itself back together in
a "big
crunch," or whether the galaxies would sail ever more slowly outward forever.
It was to measure that rate of slowing of this outward flight, and thus find the
long-sought
and elusive answer to the cosmic question, that two teams of astronomers started
competing projects in the 1990's using distant exploding stars, supernovas, as
cosmic
beacons.
In 1998 the two teams announced that instead of the expected slowing, the galaxies
actually seem to have speeded up over the last five or six billion years, as if
some "dark
energy" was pushing them outward.
"It's definitely the strangest experimental finding since I've been in physics,"
Dr. Witten
said. "People find it difficult to accept. I've stopped expecting that the finding
will be
proved wrong, but it's an extremely uncomfortable result."
To astronomers this dark energy bears a haunting resemblance to an idea that Albert
Einstein had back in 1917 and then abandoned, later calling it his biggest blunder.
In that
year he inserted a mathematical fudge factor that came to be known as the
cosmological
constant into his equations of general relativity in order to stabilize the
universe against
collapse; Einstein's constant acted as a kind of cosmic repulsion to balance the
gravitational pull of the galaxies on one another.
38Einstein gave up the cosmological constant after the American astronomer Edwin
Hubble
discovered that the universe was expanding and thus did not need stabilizing. But
his
fudge factor refused to die. It gained a new identity with the advent of quantum
mechanics, the bizarre-sounding rules that govern the subatomic realm. According to
those rules, empty space is not empty, but rather foaming with energy. Inserted
into
Einstein's equations, this energy would act like a cosmological constant, and try
to blow
the universe apart.
According to astronomers the recently discovered dark energy now accounts for about
two-thirds of the mass of the universe. But is this Einstein's old fudge factor,
the
cosmological constant, come home to roost -- in which case the universe will
accelerate
eternally? Or is the presumed acceleration only temporary, driven by one of the
many
mysterious force fields, dubbed quintessence, allowed by various theories of high
energy
physics?
Or is the acceleration even real?
"It's important to find out if the cosmological constant is really constant," said
Dr. Witten.
Because the repulsive force resides in space itself, as the universe grows, the
push from
dark energy grows as well. "If dark energy is the cosmological constant then it is
a
property of the vacuum that will always be with us, getting more powerful as the
universe
gets bigger and the universe will expand forever," explained Dr. Adam Riess of the
Space
Telescope Science Institute in Baltimore. But if the dark energy is some form of
quintessence, "then there may be more such fields which arise in the future,
possibly of
the opposite sign, and then all bets are off for the future of the universe."
Dr. Krauss said, "The good news is that we can't prove that this is the worst of
all
possible universes."
The Long Goodbye
It might seem strange or presumptuous for astronomers to try to describe events all
the
way to the end of time when physicists are still groping for a "theory of
everything." But
to Dr. Krauss, this is testimony to the power of ordinary physics. "We can still
put
ultimate limits on things without even knowing the ultimate theory," he said. "We
can put
limits on things based on ordinary physics."
Dr. Dyson said his venture into eschatology was inspired partly by a 1977 paper on
the
future of an ever expanding universe by Dr. J. N. Islam, now at the University of
Chittagong in Bangladesh, in The Quarterly Journal of the Royal Astronomical
Society.
Dr. Dyson was also motivated, he wrote in his paper, to provide a counterpoint to a
famously dour statement by Dr. Steven Weinberg, who wrote in his book "The First
Three Minutes," "The more the universe seems comprehensible, the more it also seems
pointless."
39Dr. Dyson wrote, "If Weinberg is speaking for the 20th century, I prefer the
18th."
If the present trend of acceleration continues this is the forecast:
In about two billion years Earth will become uninhabitable as a gradually warming
Sun
produces a runaway greenhouse effect. In five billion years the Sun will swell up
and die,
burning the Earth to a crisp in the process. At about the same time the Milky Way
will
collide with its twin the Andromeda galaxy, now about two million light-years away
and
closing fast, spewing stars, gas and planets across intergalactic space.
Any civilization that managed to survive these events would face a future of
increasing
ignorance and darkness as the accelerating cosmic expansion rushes most of the
universe
away from us. "Our ability to know about the universe will decrease with time,"
said Dr.
Krauss. "The longer you wait, the less you see, the opposite of what we always
thought."
As he explains it, the disappearance of the universe is a gradual process. The
faster a
galaxy flies away from us, the dimmer and dimmer it will appear, as its light is
"redshifted" to lower frequencies and energies, the way a police siren sounds lower
when
it is receding. When it reaches the speed of light, the galaxy will appear to
"freeze," like a
dancer caught in midair in a photograph, in accordance to Einstein's theory of
relativity,
and we will never see it get older, said Dr. Abraham Loeb, an astronomer at
Harvard.
Rather it will simply seem dimmer. The farther away an object is in the sky, he
said, the
younger it will appear as it fades out of sight. "There is a finite amount of
information we
can collect from the universe," Dr. Loeb said. About 150 billion years from now
almost
all of the galaxies in the universe will be receding fast enough to be invisible
from the
Milky Way. The exceptions will be galaxies that are gravitationally bound to the
cloud of
galaxies, known as the Local Group, to which the Milky Way belongs. Within this
cloud,
life would look much the same at first. There would be galaxies in the sky. "When
you
look at the night the stars will still be there," said Dr. Krauss. "To the
astronomer who
wants to see beyond, the sky will be sadly empty. Lovers won't be disturbed --
scientists
will be."
But about 100 trillion years from now, when the interstellar gas and dust from
which new
stars condense is finally used up, new stars will cease to be born. From that time
on, the
sky will grow darker and darker. The galaxies themselves, astronomers say, will
collapse
in black holes within about 1030 years.
But even a black hole is not forever, as Dr. Stephen Hawking, the Cambridge
University
physicist and best-selling author, showed in path-breaking calculations back in
1973.
Applying the principles of quantum mechanics to these dread-sounding objects, Dr.
Hawking discovered that a black hole's surface, its so-called event horizon, would
fluctuate and exude energy in the form of random bursts of particles and radiation,
growing hotter and hotter until the black hole eventually exploded and vanished.
Black holes the mass of the sun would take 1064 years to explode. For black holes
the
mass of a galaxy those fireworks would light up space-time 1098 years from now.
40Against the Fall of Night
Will there be anything or anyone around to see these quantum fireworks?
Dr. Dyson argued in his 1979 paper that life and intelligence could survive the
desert of
darkness and cold in a universe that was expanding infinitely but ever more slowly
by
adopting ever slower and cooler forms of existence. Intelligence, could reside, for
example, in the pattern of electrically charged dust grains in an interstellar
cloud, a
situation described in the 1957 science fiction novel "The Black Cloud," by the
British
astronomer Sir Fred Hoyle, who died in August.
As an organism like the black cloud cooled, he argued, it would think more slowly,
but it
would always metabolize energy even more slowly, so its appetite would always be
less
than its output. In fact, Dr. Dyson concluded, by making the amount of energy
expended
per thought smaller and smaller the cloud could have an infinite number of thoughts
while consuming only a finite amount of energy.
But there was a hitch. Even just thinking requires energy and generates heat, which
is
why computers have fans. Dr. Dyson suggested that creatures would have to stop
thinking and hibernate periodically to radiate away their heat.
In an accelerating universe, however, there is an additional source of heat that
cannot be
gotten rid of. The same calculations that predict black holes should explode also
predict
that in an accelerating universe space should be filled with so-called Hawking
radiation.
In effect, the horizon -- the farthest distance we can see -- looks mathematically
like the
surface of a black hole. The amount of this radiation is expected to be incredibly
small --
corresponding to a fraction of a billionth of a billionth of a billionth of a
degree above
absolute zero, but that is enough to doom sentient life.
"The Hawking radiation kills us because it gives a minimum temperature below which
you cannot cool anything," said Dr. Krauss. Once an organism cools to that
temperature,
he explained, it would dissipate energy at some fixed rate. "Since there is a
finite total
energy, this means a finite lifetime."
Infinity on Trial
Although Dr. Dyson agrees with this gloomy view of life in an accelerating
universe, he
and Dr. Krauss and Dr. Starkman are still arguing about whether life is also doomed
in a
universe that is not accelerating, but just expanding and getting slower and
colder.
Quantum theory, the Case Western authors point out, limits how finely the energy
for
new thoughts can be shaved. The theory decrees that energy is emitted and absorbed
in
tiny indivisible lumps called "quanta." Any computation must spend at least this
much
energy out of a limited supply. Each new thought is a step down an energy ladder
with a
41finite number of steps. "So you can only have a finite number of thoughts," said
Dr.
Krauss.
"If you want to stare at your navel and not think any new thoughts, you won't
dissipate
energy, " he explained. But that would be a boring way to spend eternity. If life
is to
involve more than the eternal reshuffling of the same data, he and Dr. Starkman
say, it
cannot be eternal.
Dr. Dyson, however, says this argument applies only to so-called digital life, in
which
there is a fixed number of quantum states. Creatures like the black cloud, which
could
grow along with the universe, he said, would have an increasing number of quantum
states, and so there would always be more rungs of the ladder to step down. So the
bottom need never be reached and life and thought could go on indefinitely.
But nobody knows whether such a life form can exist, said Dr. Krauss.
Compared with the sight of the World Trade Center towers collapsing or the plight
of a
sick child, this future extinction may seem a remote concern. Dr. Allan Sandage, an
astronomer at Carnegie Observatories in Pasadena, Calif., who has spent his life
investigating the expansion and fate of the universe, said: "Life on this earth is
going to
vanish in 4.5 billion years. I wouldn't get hung up on the fact that the lights are
all going
out in 30 billion years."
Dr. Dyson said he was still an optimist. It is too soon to start panicking, he
counseled in
an e-mail message. The observations could be wrong.
"At present all possibilities are open," he wrote. "The recent observations are
important,
not because they answer the big questions about the history of the universe, but
because
they give us new tools with which to explore the history."
Even in an accelerating universe, Dr. Dyson said, humans or their descendants might
one
day be able to rearrange the galaxies and save more of them from disappearing.
Another
glimmer of hope comes from the deadly and chilling Hawking radiation itself, said
Dr.
Raphael Bousso, from the Institute of Theoretical Physics at the University of
California
at Santa Barbara. Since that radiation is produced by unpredictable quantum
fluctuations,
he pointed out, if you wait long enough anything can appear in it, even a new
universe.
"Sooner or later one of those quantum fluctuations will look like a Big Bang," he
said.
In that case there is the possibility of a future, if not for us, at least for
something or
somebody. In the fullness of time, after all, physics teaches that the improbable
and even
the seemingly impossible can become the inevitable. Nature is not done with us yet,
nor,
as Dr. Dyson indicates, are we necessarily done with nature.
We all die, and it is up to us to decide who and what to love, but, as Dr. Weinberg
pointed out in a recent article in The New York Review of Books, there is a certain
nobility in that prospect.
42"Though aware that there is nothing in the universe that suggests any purpose for
ure brings about an increase of the thick-
ness and a shortening of the surface, while, on the
other hand, tension leads to splitting of the continen-
tal blocks. The individual stages of perceived as
mountain formation comprised continual processes
of splitting and compression, whereby the original
Sialic crust (for which Wegener assumed a thickness
of about 30–35 km) gradually decreased in surface
area, split into separate pieces, and increased in thick-
ness. Along with the movement of the continental
blocks, a hypothesized universal ocean (‘Panthalassa’)
began to divide into a shallow sea and a deep sea.
Volcanism, for Wegener, was mainly related to the
continental ‘fronts’. Areas where tension prevailed,
such as the Atlantic Ocean, and also opening faults,
seemed to be relatively poor in volcanoes as compared
with areas such as the Pacific Ocean, where pressure
was increasing. The fronts of moving blocks made
conditions more favorable to volcanism than did the
‘backs’. Nevertheless, Wegener wondered whether the
mid-Atlantic ridge might be considered as a zone
where, with the continuing expansion of the Atlantic,
the floor was continuously breaking up, making room
for fresh, relatively fluid and high-temperature Sima
from below! Moreover, increased volcanic activity in
some periods of Earth history might be due to large
displacements (as, for instance, during the Tertiary).
Trench faults (Grabenbru che), i.e., rift valleys,
acquired new meaning as representing the beginnings
of new continental separations. Gravity measure-
ments had shown that beneath such lines lay material
of greater density, compared to that on either side.
Therefore, these lines could be seen as incipient fis-
sures within the continental blocks (into which the
denser Sima was rising according to the principle of
isostasy). The best examples of such separations were
provided by the East African trenches and their con-
tinuation through the Red Sea. At the majority of the
trenches, the measurable mass deficit was not com-
pensated by greater density of the matter beneath it.
Thus, the trenches must be youthful disruptions of a
continental block.
Wegener’s theory of mountain formation was fur-
ther supported by the fact that the folding of the
Andes seems to have been essentially simultaneous
to the opening of the Atlantic Ocean. The American
blocks, during their westward drifting, had encoun-
tered resistance at the presumably very old and rela-
tively rigid floor of the Pacific Ocean. Thus, the
extended shelf, with its mighty sediments, forming
the western border of the continental block, was
compressed to a range of fold mountains. For the
Tertiary folds of the Himalayas, Wegener assumed
that lower India had formed an extended peninsula
prior to compression, the southern end of which lay
next to that of South Africa. The folds had been
produced by ‘impact’ of the Indian subcontinent and
the main mass of Asia.
Geological and Palaeontological
Evidence
The palaeontological evidence indicating a former
connection between the organic components of differ-
ent continents had already given rise to the doctrine
of former land bridges. Among the most striking
findings were the distributions of the Glossopteris
flora on the southern continents and the occurrence
of Mesosaurus at the turn of the Permian and the
Carboniferous exclusively in south-eastern South
America and the western parts of Africa; both of
these discoveries suggested a former connection of
the two continents. Using these relationships also
allowed calculations of when the continents were
separated (either by horizontal displacements or by
sinking of the land bridges). South America and Africa
had been connected during the Mesozoic, but were
separated at the end of the Eocene or Early Oligocene.
The connection between Europe and North America
seemed to have been maintained during the older
Tertiary period, but separation occurred in the
Miocene, although it might have continued in the far
north (over Scandinavia and Greenland) into the
Pleistocene. The connection of Lower India with
southern Africa, which Wegener had postulated
based on his ideas on the formation of the Himalayan
range, was also confirmed by palaeontological evi-
dence. Zoogeographers had long assumed a former
elongated Indian–Madagascan peninsula (called
‘Lemuria’), separated from the African block by the
Mozambique Channel.
The zoogeographic concept of Lemuria had given
rise to Suess’ notion of a great southern contin-
ent, Gondwana, comprising parts of South America,
Africa, Lower India, Australia, and Antarctica.
Assuming the unchanged positions of its present-day250 FAMOUS GEOLOGISTS/Wegener
relics, however, required ascribing a huge extent to
this continent. Wegener, by contrast, proposed a
much reduced primeval continent, Pangaea. In the
Permian, i.e., until some 300 Ma ago, all the contin-
ents were supposedly joined in one land mass
extending from pole to pole. During the Triassic,
about 200 Ma ago, Pangaea began to break up and
the newly emerging continents started moving into
their current positions. In the Jurassic, there were
few remaining connections except at the northern
and southern ends. Just as northern Europe and
North America remained connected until the older
Tertiary period, a connection of the southern con-
tinents seems to have persisted, running from the
southern coast of Australia over Antarctica to South
America. Later, the Antarctic block, like the
South American block in the Tertiary, moved over
from South Africa towards the side of the Pacific
Ocean. Only in the Quaternary period, then, did the
Australian block become detached (Figure 4).
For geological and tectonic evidence, Wegener
referred particularly to Suess’ magnum opus, pub-
lished in three volumes during 1885–1909, Das
Antlitz der Erde (The Face of the Earth). Considering
the tectonic relations, Europe/Africa and both Amer-
icas seemed to represent the edges of an immense
expanded fissure. In the north, for instance, the Green-
land massif was matched by Scandinavia, both consist-
ing of gneiss, and the less mountainous North
America corresponded to the likewise less mountain-
ous Europe. The most striking example, however, was
the Carboniferous mountain range, called the Armor-
ican mountains (Suess’ ‘transatlantic Altaides’),
which made the coalfields of North America appear
to be the direct continuation of the European ones.
Wegener’s theory of mountain formation was also
confirmed by remarkable differences between the
Atlantic and the Pacific hemispheres, such as the
distinction between Pacific and Atlantic types of
coasts (marginal chains and ocean trenches in front
Figure 4 Wegener’s reconstruction of the separation of the continents from the
primeval Pangaea, from his 1926 paper Pala ogeo
graphische Darstellung der Theorie der Kontinentalverschiebungen , showing the
relative positions of the continents during the Upper
Carboniferous (Jung Karbon), Eocene (Eozan), and Lower Quaternary (Alt Quartar) (in
two different projections). Cross hatching
represents deep seas, dotted regions represent shallow seas; rivers, recent
coastlines, and outlines are shown only for orientation.FAMOUS GEOLOGISTS/Wegener
251
of the Pacific coasts, as contrasted to the wild, irregu-
lar ‘ria’ Atlantic coastlines). There were also differ-
ences in the volcanic lavas of the two hemispheres, as
emphasized by the Vienna petrographer Friedrich
Becke (1855–1931) and others. The Atlantic lavas
contained a greater proportion of sodium, whereas
calcium and magnesium prevailed in the Pacific
lavas. Such differences were intelligible according to
the assumptions of continental movements. The
opening of the Atlantic was matched by the general
pressing of the continents against the region of the
Pacific Ocean: pressure and compression prevailed
at the coasts of the latter whereas tension and
splitting occurred at the latter.
Palaeoclimatology
Traces of glaciation during the Permian (ground mo-
raines lying on scratched bedrock) were to be found
on the southern continents, e.g., in East India and
Australia. If the present-day arrangement of the land
masses had prevailed at that time, this ‘Permian ice
age’ would have required an icecap of seemingly im-
possible size. And the north pole would have been in
Mexico, where no trace of glaciation during that
period was recorded. Following the idea of horizontal
displacements, however, all regions subjected to
glaciation came together concentric to the southern
margin of Africa. And one had only to place the south
pole in this much reduced glaciated area to give the
Permian ice age a much more plausible form.
Wegener had discussed these palaeoclimatological
features since 1912. In 1924, he gave a detailed
description of the climatological changes from the
Carboniferous through to recent times, following the
traces of glaciations, swamps, and deserts, i.e., mo-
raines, coal, salt, and gypsum, throughout Earth’s
history (Figure 5). In reconstructing the respective
polar shifts, Wegener emphasized that they obviously
took place along with the great displacements of the
continental blocks. In particular, there was temporal
coincidence of the best confirmed polar shift, in the
Tertiary, and the opening of the Atlantic (Figure 6).
Movement of the poles since the Pleistocene might also
be related to the final separations of the continents in
the north and the south.
Motive Forces
Wegener was very cautious about the forces that might
have caused continental displacements. First, it was
necessary to demonstrate the reality and the manner of
the displacements before indulging in the hope of find-
ing their cause. Nevertheless, he tentatively suggested
two candidates: centrifugal forces caused by the rota-
tion of Earth and tidal-type waves within Earth, gen-
erated by the gravitational pull of the sun and the
moon. In the 1929 revision of Wegener’s theory in
Figure 5 Wegener thought continental drift was the key to the climatic changes
during Earth’s history. This map, published in the
1924 book by Koppen and Wegener, Die Klimate der geologischen Vorzeit , shows
traces of glaciation, swamps, and deserts for the
Carboniferous. E, Traces of glaciation; K, coal; S, salt; G, gypsum; W, desert
sandstone. Dotted regions indicate arid areas, dashed
lines indicate the positions (i.e., the pathways) of the poles, and the bold curved
line indicates the respective position of the equator.252 FAMOUS GEOLOGISTS/Wegener
Figure 6 Map published in the 1924 book by Koppen and Wegener, Die Klimate der
geologischen Vorzeit , showing polar shifts (dashed
lines) from the Carboniferous to recent, related to the African table (left, south
pole; right, north pole). Bold lines outline the continental
blocks; hatched lines represent the Carboniferous (Karbon) period. Perm, Permian;
Jura, Jurassic; Trias, Triassic; Kreide, Cret
aceous; Eozan, Eocene; Miozan, Miocene; Beginn des Quartar, beginning of the
Quaternary.
Die Klimate, he also mentioned convection currents
within the Sima; these had been first discussed as a
cause of mountain formation by the Vienna geologist
Otto Ampferer (1875–1947) in 1906.
Wegener also endeavoured to calculate the recent
velocity of the relative motion of the continents, thou-
gh he was well aware that these values must be
quite uncertain. In his 1912 paper, comparing various
longitude determinations for Greenland, he had
deduced an increase of the distance to Europe of
11 m year 1 . Referring to the lengths of transatlantic
cables, he suggested that North America was drifting
away from Europe at about 4 m year 1 .
From Continental Drift to
Plate Tectonics
The theory of continental drift was long rejected by
the majority of geologists. Among Wegener’s few
followers were the South African Alexander Du Toit
(1878–1948), for whom continental drift provided
the best explanation of the close similarities between
the strata and fossils of Africa and South America,
and the Swiss geologist E mile Argand (1879–1940),
who saw continental collisions as the only means of
producing the folded and buckled strata he had ob-
served in the Alps (see Famous Geologists: Du Toit).
Nevertheless, Wegener’s explanation of the Permo-
Carboniferous ice age impressed even his critics.
Wegener’s reputation as a meteorologist and a
polar explorer contributed to keeping his theory
alive. His work was immediately remembered when,
around 1960, surprising data were obtained from the
ocean floor: palaeomagnetic patterns alongside the
mid-ocean ridges clearly suggested the spreading of
the seafloor. Within about two decades, Wegener’s
principle of horizontal displacements of parts of
Earth’s crust became almost universally accepted, al-
though, ironically, the process still lacked a consensus
as to its causes, though convection currents in the
internal mantle are most commonly advocated.
It should be noted that Wegener’s original concept
differed from modern plate tectonics in essential
points, particularly with regard to the Sial and the
Sima. According to modern theory, the (Sialic) con-
tinents do not ‘plough’ through the (oceanic) Sima.
Instead, both continents and ocean floor are regarded
as forming solid plates, ‘floating’ on the astheno-
sphere, which, due to tremendous heat and pressure,
behaves like an extremely viscous liquid (as Wegener
had thought the Sima did). Therefore, the older
term ‘continental drift’, still often used today, is not
quite appropriate for the modern concept. Notwith-
standing these differences, Wegener’s basic ideas
remain sound, and the lines of evidence that he
used to support his theory are still valid. He first
envisaged a dynamic Earth, connecting its major
features and various geological processes – continentalFLUID INCLUSIONS 253
movements, folded mountain ranges, rift systems,
earthquakes, volcanism, ocean transgressions, pal-
aeoclimatological changes, etc. – on a global scale. In
this sense, Wegener’s theory was a true forerunner of
plate tectonics.
See Also
Africa: Rift Valley. Famous Geologists: Du Toit; Suess.
Gondwanaland and Gondwana. History of Geology
From 1900 To 1962. History of Geology Since 1962.
Palaeoclimates. Pangaea. Plate Tectonics. Tectonics:
Mid-Ocean Ridges; Mountain Building and Orogeny.
Further Reading
Carozzi AV (1985) The reaction of continental Europe
to Wegener’s theory of continental drift. Earth Sciences
History 4: 122 137.
Fritscher B (2002) Alfred Wegener’s ‘The origin of contin
ents, 1912’. Episodes 25: 100 106.
Jacoby WR (2001) Translation of ‘Die Entstehung der Kon
tinente, Dr Alfred Wegener, Petermann’s Geographische
Mitteilungen, 58 (1912)’. Journal of Geodynamics 32:
29 63.
Ko ppen V and Wegener A (1924) Die Klimate der geolo
gischen Vorzeit. Berlin: Borntra ger.
Lu decke C (1994) Stratigraphische Methode der Rekon
struktion von Expeditionsergebnissen am Beispiel des
Todes von Alfred Wegener wa hrend der Gro nlandexpedi
tion (1930 31). In: Fritscher B and Brey G (eds.) Cosmo
graphica et Geographica: Festschrift fu r Heribert M.
Nobis zum 70. Geburtstag, Algorismus, vol. 13,
pp. 347 367. Munich: Institut fu r Geschichte der
Naturwissenschaften.
Oreskes N (1999) The Rejection of Continental Drift:
Theory and Method in American Earth Science. New
York and Oxford: Oxford University Press.
Runcorn SK (ed.) (1966) Continental Drift. New York and
London: Academic Press.
Schwarzbach M (1986) Alfred Wegener: The Father
of Continental Drift. Madison. WI: Science Tech
Publications.
Sengo r AMC (1991) Timing of orogenic events: a persistent
geological controversy. In: Mu ller DW, McKenzie JA,
and Weissert H (eds.) Controversies in Modern Geology:
Evolution of Geological Theories in Sedimentology,
Earth History and Tectonics, pp. 403 473. London:
Academic Press.
Wegener A (1912) Die Entstehung der Kontinente.
Petermann’s Mitteilungen aus Justus Perthes’ Geogra
phischer Anstalt 58: 185 195, 253 256, 305 309.
Wegener A (1926) Pala ogeographische Darstellung der
Theorie der Kontinentalverschiebungen. In: Dacque E
(ed.) Pala ogeographie, pp. 171 189. Leipzig and Wien:
F Deuticke.
Wegener A (1971) The Origin of Continents and Oceans.
(Translation from the 4th revised German edition by
J Biram, with an introduction by BC King.) London:
Methuen.
Wegener A (1980) Die Entstehung der Kontinente und
Ozeane. (Reprint of the 1st and 4th editions, edited by
A Vogel.) Braunschweig: Vieweg.
Wegener E (1960) Alfred Wegener: Tagebu cher, Briefe,
Erinnerungen. Wiesbaden: Brockhaus.
Wutzke U (1998) Kommentiertes Verzeichnis der schriftli
chen Dokumente seines Lebens und Wirkens, Berichte
zur Polarforschung 288. Bremerhaven: Alfred Wegener
Institut fu r Polar und Meeresforschung.
FLUID INCLUSIONS
A H Rankin, Kingston University,
Kingston-upon-Thames, UK
ß 2005, Elsevier Ltd. All Rights Reserved.
Introduction
Fluid inclusions are small droplets of fluid that have
been trapped within crystals either during primary
growth from solution or at some later stage, usually
as a result of recrystallization along healed micro-
fractures. They are ubiquitous in both naturally
occurring minerals and in laboratory-grown crystals.
To the chemist or materials scientist, these gross
defects cause endless obstacles in their quest to grow
near-perfect crystals. However, to the geologist, they
provide a unique fossil record of the various
fluids responsible for the formation and evolution of
rocks and minerals throughout the history of the
Earth.
Despite their small size (usually less than 20 mm),
their chemical composition and physical properties
can be readily determined, and the data may be used
to estimate the temperatures, pressures, and physico-
chemical nature of the fluid at the time of trapping.
This information has made an immense contribution
to the development of modern theories of ore genesis,
petrogenesis, diagenesis, and petroleum migration
and accumulation, and to our understanding of the
importance of the fluid phase in a wide range of
geological processes.254 FLUID INCLUSIONS
Occurrence and General
Characteristics
Formation and Genetic Classification of Fluid
Inclusions
Small changes in the chemical or physical properties
of fluids near to a growing crystal face can lead to
perturbations in the stability of crystal growth and the
development of gross defects, manifested as embay-
ments, along crystal faces. These embayments will
seal over during a period of greater stability, trapping
a portion of fluids to form ‘primary’ (P) fluid inclu-
sions. In many instances, the trapped fluid will be
‘homogeneous’ at the time of trapping. In others,
where immiscible fluids are present or where mechan-
ical entrapment of other coexisting crystalline phases
has occurred, trapping will be ‘heterogeneous’.
At some stage after primary growth, ‘secondary’ (S)
fluid inclusions can form from later fluids, particularly
as a result of recrystallization along microfractures.
The chemical and physical properties of these inclu-
sions may be very different from those of the earlier
mineral-forming fluids. However, if fracturing and
rehealing take place during primary growth, the fluids
may be indistinguishable, and the terms ‘pseudose-
condary’ or ‘primary–secondary’ (PS) appropriately
describe such inclusions. A schematic representation
of this genetic classification of inclusions is shown in
Figure 1.
For most geological applications, it is necessary to
establish whether the inclusions are primary, second-
ary, or pseudosecondary, and also whether heteroge-
neous trapping has occurred. Heterogeneous trapping
may be recognized by the variable proportions of
liquids and solids in a single group or generation of
inclusions. Various criteria may be used to distinguish
between P, PS, and S inclusions, but these may be
difficult to apply and it may be difficult to identify
primary inclusions in many samples.
Figure 1 Schematic representation of the distribution of pri
mary (P), secondary (S), and pseudosecondary (PS) fluid inclu
sions in a quartz crystal. Modified from Rankin AH (1989) Fluid
inclusions. Geology Today 5: 21 24.
in turbid or translucent minerals, such as feldspar.
Quartz is usually the preferred host.
Size and Shape of Inclusions
Choice of Material for Study
The successful application of fluid inclusion studies
depends partly on serendipity and partly on the type
and quality of material available for study. Due to
their small size, observations on fluid inclusions are
carried out under a microscope using polished wafers
around 1–2 mm thick. In most cases, clear, transpar-
ent minerals are needed, but it is also possible to study
inclusions in some deeply coloured, semi-transparent
minerals in very thin (<50 mm) polished sections.
Care must be taken with soft, easily cleaved minerals,
such as calcite and fluorite, because of the possibility
of leakage during sample preparation or analysis.
Fluid inclusions are particularly difficult to observe
Fluid inclusions seldom exceed 1 mm in size; most are
less than 20 mm, and those greater than 1 cm are
exceptionally rare and usually regarded as museum
specimens. Fluid inclusions display a variety of
shapes. They may be flattened and irregular, rounded,
or regular with three-dimensional ‘negative crystal’
shapes mimicking the crystal symmetry of the host
crystal (Figure 2).
Phases Present at Room Temperature
Fluid inclusions contain varying proportions of liquid
(L), solid (S) and gas (G) depending on the compos-
ition (X), temperature (T), pressure (P), and volume
(V) of the enclosed fluid at the time of entrapment.
humanity," he wrote, "one way that we can find a purpose is to study the universe
by the
methods of science, without consoling ourselves with fairy tal
Niels Bohr, a Danish physicist and leader of this revolution, once said that a
person who
was not shocked by quantum theory did not understand it.
This week, some 700 physicists and historians are gathering in Berlin, where Planck
started it all 100 years ago, to celebrate a theory whose meaning they still do not
understand but that is the foundation of modern science. Quantum effects are now
7invoked to explain everything from the periodic table of the elements to the
existence of
the universe itself.
Fortunes have been made on quantum weirdness, as it is sometimes called.
Transistors
and computer chips and lasers run on it. So do CAT scans and PET scans and M.R.I.
machines. Some computer scientists call it the future of computing, while some
physicists
say that computing is the future of quantum theory.
"If everything we understand about the atom stopped working," said Leon Lederman,
former director of the Fermi National Accelerator Laboratory, "the G.N.P. would go
to
zero."
The revolution had an inauspicious start. Planck first regarded the quantum as a
bookkeeping device with no physical meaning. In 1905, Albert Einstein, then a
patent
clerk in Switzerland, took it more seriously. He pointed out that light itself
behaved in
some respects as if it were composed of little energy bundles he called
lichtquanten. (A
few months later Einstein invented relativity.)
He spent the next decade wondering how to reconcile these quanta with the
traditional
electromagnetic wave theory of light. "On quantum theory I use up more brain grease
than on relativity," he told a friend.
The next great quantum step was taken by Bohr. In 1913, he set forth a model of the
atom
as a miniature solar system in which the electrons were limited to specific orbits
around
the nucleus. The model explained why atoms did not just collapse -- the lowest
orbit was
still some slight distance from the nucleus. It also explained why different
elements
emitted light at characteristic wavelengths -- the orbits were like rungs on a
ladder and
those wavelengths corresponded to the energy released or absorbed by an electron
when
it jumped between rungs.
But it did not explain why only some orbits were permitted, or where the electron
was
when it jumped between orbits. Einstein praised Bohr's theory as "musicality in the
sphere of thought," but told him later, "If all this is true, then it means the end
of
physics."
While Bohr's theory worked for hydrogen, the simplest atom, it bogged down when
theorists tried to calculate the spectrum of bigger atoms. "The whole system of
concepts
of physics must be reconstructed from the ground up," Max Born, a physicist at
Gottingen University, wrote in 1923. He termed the as-yet-unborn new physics
"quantum
mechanics."
Boy's Mechanics
The new physics was born in a paroxysm of debate and discovery from 1925 to 1928
that
has been called the second scientific revolution. Wolfgang Pauli, one of its
ringleaders,
called it "boy's mechanics," because many of the physicists, including himself,
then 25,
8Werner Heisenberg, 24, Paul Dirac, 23, Enrico Fermi, 23, and Pascual Jordan, 23,
were
so young when it began.
Bohr, who turned 40 in 1925, was their father-confessor and philosopher king. His
new
institute for theoretical physics in Copenhagen became the center of European
science.
The decisive moment came in the fall of 1925 when Heisenberg, who had just returned
to
Gottingen University after a year in Copenhagen, suggested that physicists stop
trying to
visualize the inside of the atom and instead base physics exclusively on what can
be seen
and measured. In his "matrix mechanics," various properties of subatomic particles
could
be computed -- but, disturbingly, the answers depended on the order of the
calculations.
In fact, according to the uncertainty principle, which Heisenberg enunciated two
years
later, it was impossible to know both the position and velocity of a particle at
once. The
act of measuring one necessarily disturbed the other.
Physicists uncomfortable with Heisenberg's abstract mathematics took up with a
friendlier version of quantum mechanics based on the familiar mathematics of waves.
In
1923, the Frenchman Louis de Broglie had asked in his doctoral thesis, if light
could be a
particle, then why couldn't particles be waves?
BALTIMORE, April 5 -- A gasp went through the auditorium of the Space Telescope
Science Institute on Wednesday when Dr. Adam Riess, a young astronomer from the
institute, put the last mark on his so-called Hubble diagram, a plot of the
brightness and
speed of distant objects that astronomers use to divine the history of the
universe.
The Darth Vaders of astronomy had gathered here to take stock of their expanding
and
increasingly dark realm. Once upon a time astronomy was about what could be seen in
the sky, about jewel-like lights that moved in eternally recurring patterns and the
soft
glow of galaxies and comets.
Now it is about what cannot be seen. In the last few decades astronomers have had
to
confront the possibility that stars and galaxies -- not to mention the creatures
that inhabit
them -- are barely more than flecks of froth on a stormy sea of dark matter.
Now Dr. Riess was presenting his colleagues with evidence, based on observations of
a
star that exploded 11 billion years ago, that the universe -- dark matter and all
-- was
being blown apart under the influence of a mysterious antigravitational force known
only
as "dark energy."
"We are doing astronomy of the invisible," admitted Dr. Mario Livio, a theorist at
the
Space Telescope institute, who had organized the meeting, called "The Dark
Universe:
Matter, Energy, and Gravity" last fall.
As it turned out, the meeting coincided with a NASA news conference announcing the
breakthrough discovery by Dr. Riess and his colleagues and thus was dominated by
discussions of new telescopes in space and new dimensions in the universe as
astronomers grappled with the meaning of dark energy and how to take its measure.
Now physicists, some of whom have been reluctant to take acceleration of the
universe
seriously, will have to explain what this dark energy is. "Those numbers are
alarming,
and apparently true," said Dr. Michael Dine, a theoretical physicist from the
University of
California at Santa Cruz. He described his colleagues as now working "frantically"
to
find an explanation.
19On one level, the universe, with all of its dark baggage, seems to make sense.
The total
amount of matter and energy seems to be just enough to guarantee that the large-
scale
geometry of space-time is "flat," or Euclidean, a result that cosmologists have
long
considered to be the most desirable and aesthetic. On the other hand, the detailed
breakdown of the constituents of the cosmos is, as Dr. Livio says, "ugly" -- 65
percent
dark matter, 30 percent dark matter of unknown nature and only 5 percent stars, gas
and
dust.
"We live in a preposterous universe," said Dr. Michael Turner, an astrophysicist at
the
University of Chicago. "Dark energy. Who ordered that?"
Of course, it was Einstein who originally ordered dark energy when he inserted a
fudge
factor called the cosmological constant into his gravitational equations describing
the
universe. Lambda, as it is known, after the Greek letter, represented a sort of
cosmic
repulsion associated with space itself that kept the cosmos from collapsing of its
own
weight. Einstein abandoned the cosmological constant when it was discovered that
the
universe was expanding, and resisted efforts to bring it back, once referring to it
as his
biggest blunder.
But he couldn't keep it out forever. In 1998 two competing teams of astronomers
trying to
measure how the expansion of the universe was slowing down because of cosmic
gravity,
found that the universe was actually speeding up, as if the galaxies were being
pushed
apart by a force -- dubbed, in the spirit of the times, "dark energy."
"This was a very strange result," recalled Dr. Riess, who was a member of one of
the
teams. "It was the opposite of what we thought we were doing." Was this Einstein's
old
cosmological constant, something even weirder or a mistake?
The effect had showed up as an unexpected dimness on the part of certain exploding
stars
known as supernovae that the astronomers were using as so-called standard candles,
objects whose distance could be gauged from their apparent brightness. The
astronomers
deduced that these stars were farther away than they should have been in an evenly
expanding universe, and that therefore the expansion was actually accelerating.
But dust or chemical changes over the eons in the stars could also have dimmed the
supernovae. The cleanest test of dark energy and the acceleration hypothesis, Dr.
Riess
explained, would be to find supernovae even farther out and back into the past,
halfway
or more back to the Big Bang itself. Because it is space itself that provides the
repulsive
push, according to Einstein's equations, that push should start out small when the
universe is small and grow as the universe expands. Cosmic acceleration would only
kick
in when lambda's push got big enough to dominate the gravity of ordinary matter and
energy in the universe, about five or six billion years ago. Before then the
universe would
have been slowing down, like a Mark McGwire blast that has not yet reached the top
of
its trajectory, and a supernova glimpsed at that great distance would appear
relatively
brighter than it should. If dust or chemical evolution were responsible, such
distant stars
should appear relatively even dimmer.
20By chance the Hubble Space Telescope had observed a supernova in late 1997 and
early
1998 that proved to be 11 billion light-years away -- the most distant yet seen. On
Dr.
Riess's Hubble diagram it appeared twice as bright as it should.
"Extraordinary claims require extraordinary evidence -- I hope the I.R.S. doesn't
say that
to you," Dr. Riess told his audience, but, he concluded, "the cosmological constant
looks
good for this supernova."
Dr. Livio said, "A year ago probably a large fraction of the people in this room
would not
have believed it."
But there were more complicated explanations, forms of dark energy other than the
cosmological constant on physicists' drawing boards, as well as the possibility
that
astronomers were still being fooled. To explicate the nature of the dark energy,
astronomers need to observe more supernovae as far back as 11 billion years ago, to
cover the time when the universe began to accelerate.
"How fast did it go from deceleration to acceleration?" asked Dr. Riess. Answering
such
questions could help astronomers determine how hard the dark energy is pushing on
the
universe compared with the predictions for the cosmological constant. A fast
turnaround,
he said, "begins to tell you there is a lot of oomph for a given amount of 'it.' "
"The cosmological constant is the benchmark oomph," he said.
To find those supernovae so far out cosmologists will have to go to space, said Dr.
Saul
Perlmutter, a physicist at the University of California's Lawrence Berkeley
National
Laboratory and a veteran dark energy hunter.
On the ground, the supernova researchers have to employ a wide network of people
and
telescopes to detect the explosions, diagnose their type and then to watch them
fade. Dr.
Perlmutter described an orbiting telescope that would perform all three functions.
The
Supernova/Acceleration Probe, or SNAP, would combine an 80-inch diameter mirror
(only about 16 percent smaller than the Hubble), a giant electronic camera with a
billion
pixels and a special spectroscope.
If all goes well, Dr. Perlmutter said, the telescope could be launched in 2008. In
three
years of operation, he estimated, SNAP could harvest about 2,000 supernovae. To
distinguish the different ideas about dark energy, observations would have to be
refined
to the level of 1 or 2 percent uncertainty .
"We're all excited," he said.
So are the physicists. Their list of suspects begins with Einstein's cosmological
constant,
but therein lies a problem. About the time that Einstein was abandoning it, quantum
mechanics -- the set of rules that govern the subatomic realm -- was establishing a
21theoretical foundation for the cosmological constant. According to quantum
theory,
empty space should be foaming with temporary particles and their cumulative energy
would outweigh the matter in the universe, including dark matter, by 120 orders of
magnitude -- that is, a factor of 10 followed by 119 zeros. At that level, the
force of the
vacuum would either have crumpled the universe or blown it apart before even an
atom
had the chance to form.
The fact that the universe is in fact puttering along rather gently suggests that
there is
something fundamental about physics and the universe that physicists still don't
know.
Dr. Steven Weinberg, the Nobel Prize-winning particle theorist at the University of
Texas, has called the cosmological constant "the bone in our throat." If the dark
energy
really is Einstein's cosmological constant, then physicists have to answer
questions like
why it is so small -- roughly comparable, in fact, to the density of matter in our
own
epoch.
Lacking an answer so far, even from string theory, the mathematically daunting
candidate
theory of everything, some theorists have resorted to a controversial and somewhat
philosophical notion called the anthropic principle, which, in effect, says that
physicists
need to factor in their own existence when they think about the universe. Out of
all the
possible universes that can be imagined, this line of thinking goes, the only ones
in which
humans can find themselves is one that is conducive to human life.
That means, Dr. Weinberg has pointed out, that the cosmological constant has to be
small
enough to allow time for galaxies and stars to condense from the primordial fog
before it
takes over and starts blowing the universe apart.
Dr. Alex Vilenkin of Tufts University in Massachusetts pointed out to the Dark
Universe
audience that the universe was at its peak in making stars about five billion or
six billion
years ago, just about the time that dark energy and the matter density would have
been
equal. Our own sun, some 4.5 billion years old, was on the tale end of that wave,
and now
here we are. "Observers are where the galaxies are," said Dr. Vilenkin. "A typical
observer will see a small cosmological constant."
Many physicists are uncomfortable with this line of reasoning, and they are seeking
the
answer in different class of theories known as quintessence, after the Greek word
for the
fifth element. Modern physics, noted Dr. Paul Steinhardt, a theorist at Princeton,
is
replete with mysterious energy fields that would exhibit negative gravity. The
trick, Dr.
Steinhardt explained, is finding a field that would act like the dark energy
without a lot of
fudging on the part of theorists.
"The observations are forcing us to do this," he said. "Dark energy is an
interesting
problem. Any solution is quite interesting."
One theory that captured the fancy of the astronomers in Baltimore was a
modification of
gravity recently proposed by three string theorists at New York University: Dr. Gia
Dvali, Dr. Gregory Gabadadze and Dr. Massimo Porrati. In string theory -- so named
22because it describes elementary particles as tiny vibrating strings -- the
ordinary world is
often envisioned as a three-dimensional island (a membrane, or "brane" in string
jargon)
floating in a 10- or 11-dimensional space. Ordinary particles like electrons and
quarks
and forces like electromagnetism are confined to three dimensions, to the brane,
but
gravity is not.
As a result, Dr. Dvali suggested that gravity could only travel so far through
conventional
space before it leaked off into the extra dimensions, thereby weakening itself. To
an
observer in the traditional three dimensions it looks as if the universe is
accelerating. The
cosmological constant, in effect, he said, is a kind of gravitational brane drain.
"Gravity
fools itself," he said. "It sees itself as a cosmological constant."
Dr. Dvali's theory was welcomed by the astronomers as a sign that string theory was
beginning to come down from its ivory tower of abstraction and make useful,
testable
predictions about the real world. (In another string contribution, Dr. Steinhardt
introduced
a new theory of the early universe, in which the Big Bang is set off by a pair of
branes
clashing together like cymbals.)
Afterward Dr. Riess and Dr. Perlmutter pressed Dr. Dvali on what they would see
when
they looked out past the crossover point where gravity began falling out of the
world;
would the transition between a decelerating universe and an accelerating one happen
more abruptly than in the case of the cosmological constant? Dr. Dvali said he
hadn't
done any calculations, but he said it was his "naïve guess" that the crossover
would
happen more smoothly than in a lambda world.
"I'd love to see this guy do some Hubble diagrams," Dr. Riess said.
Even if Dr. Dvali could be coaxed into providing a prediction, however, success in
identifying the dark energy was not guaranteed to the astronomers. Calling himself
a
spokesman for the "cranky point of view," Dr. Steinhardt pointed out that the oft-
proclaimed era of "precision cosmology" was bound to have its limits. Other
cosmological parameters, particularly the cosmic density of matter in the universe,
were
not likely to be known well enough for even SNAP to untangle the models in which
the
quintessence varied over time. Worried that the overselling of SNAP could sap
astronomers' will to come up with new ideas, he said, "We should try to make as few
pronouncements as possible."
Dr. Turner refused to be swayed from his "irrational exuberance." Appealing to the
astronomers' pride, he urged them to be ambitious. "We have a chance to do
fundamental
physics here," he said. "Let's see if we can crack this nut. Maybe we'll fall on
our faces.
Maybe cranky Paul is right.
"I still have a lot of youthful juices in my body."
IMAGINARY TIME
Before the Big Bang, There Was . . . What?
By DENNIS OVERBYE
What was God doing before he created the world? The philosopher and writer (and
later
saint) Augustine posed the question in his "Confessions" in the fourth century, and
then
came up with a strikingly modern answer: before God created the world there was no
time and thus no "before." To paraphrase Gertrude Stein, there was no "then" then.
Until recently no one could attend a lecture on astronomy and ask the modern
version of
Augustine's question -- what happened before the Big Bang? -- without receiving the
same frustrating answer, courtesy of Albert Einstein's general theory of
relativity, which
describes how matter and energy bend space and time.
If we imagine the universe shrinking backward, like a film in reverse, the density
of
matter and energy rises toward infinity as we approach the moment of origin. Smoke
pours from the computer, and space and time themselves dissolve into a quantum
"foam."
"Our rulers and our clocks break," explained Dr. Andrei Linde, a cosmologist at
Stanford
University. "To ask what is before this moment is a self-contradiction."
But lately, emboldened by progress in new theories that seek to unite Einstein's
lordly
realm with the unruly quantum rules that govern subatomic physics -- so-called
quantum
gravity -- Dr. Linde and his colleagues have begun to edge their speculations
closer and
closer to the ultimate moment and, in some cases, beyond it.
Some theorists suggest that the Big Bang was not so much a birth as a transition, a
"quantum leap" from some formless era of imaginary time, or from nothing at all.
Still
others are exploring models in which cosmic history begins with a collision with a
Inspired by de Broglie's ideas, the Austrian Erwin Schrodinger, then at the
University of
Zurich and, at 38, himself older than the wunderkind, sequestered himself in the
Swiss
resort of Arosa over the 1925 Christmas holidays with a mysterious woman friend and
came back with an equation that would become the yin to Heisenberg's yang.
In Schrodinger's equation, the electron was not a point or a table, but a
mathematical
entity called a wave function, which extended throughout space. According to Born,
this
wave represented the probability of finding the electron at some particular place.
When it
was measured, the particle was usually in the most likely place, but not guaranteed
to be,
even though the wave function itself could be calculated exactly.

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