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 THE FUNCTION OF MANGANESE IN PLANTS
WV.P. KELLEY

Historicalintroduction
Dating fromthe time of SCHEELE (i), numerousinvestigators
have noted the presenceof manganese in plants of various orders.
While small amountsonly of this elementhave been foundin most
plants, in i86o HILGARD (2) pointed out that the ash of the long-
leaf pine fromMississippi contains a relativelyhigh percentage;
and in i878 J. SCHROEDER (3) found in the ash of the Norway
spruce (Picea excels) 35.53 per cent Mn3O4,and in the ash of
the bark 4I. 23 per cent. These and otherobservations.however,
received but little attention for many years. Manganese occurs
in small amounts in practically all soils, but being an element
unessentialto growthand normal development,its absorptionby
plants was consideredto be withoutphysiologicalsignificance.
The discoveryof BERTRAND (4) in i897 of the occurrenceof
manganese in the oxidizing enzymes of plants, however,and the
subsequent findingthat small amounts of manganese salts stimu-
late the oxygen-carrying power of these catalytic agents, have
drawnattentionto thisquestionand have led to the view that after
all a physiologicalrole is probably played by this element. Since
the time of these discoveriesa large numberof experiments,with
a wide range of plants, have been made, various compounds of
manganesein water and soil culturesbeing used; and, in general,
it has been found that small amounts of manganese bring about
stimulationin growth. While small amounts oftenproduce stimu-
lation, wherevermore than a very low concentrationhas been
employed,toxicityhas resulted.
LOEW (5) and his co-workersin Japan found,furthermore, the
toxic concentrationin water cultures to be different for different
species of plants. A concentrationthat was stimulatingto rice,
for instance, proved to be toxic to barley. Likewise, SALOMONE
(6) observedin fieldexperimentsthat the applicationof such small
amounts as 4o kilogramsper hectar of manganous sulphate pro-
213] [BotanicalGazette,vol. 57
2I4 BOTANICAL GAZETTE [MARCH

duced injurious effectson wheat, while other investigatorshave

found stimulationto result from the application of much larger
amounts. In a number of instances,however,the application of
differentamounts of manganese in field culture has produced no
apparent effects.
Regarding the specific effects produced by manganese,
BERTRAND, as stated above, and a numberof other investigators,
have shown that, on the one hand, the addition of small amounts
of soluble manganese increases the oxygen-carrying power of the
oxidases, and on the other, stimulationis produced in the oxi-
dizingpower of infusionsfromplants grownunder the influenceof
manganese compounds. During recentyears an increasingimpor-
tance has been attached to the oxidizingconditionsof the soil,and,
as has been pointedout in the researchesof SULLIVAN and REID (7),
there appears to be a direct correlationbetween the growth of
plants (fertilityof the soil) and the oxidationsgoing on in the soil.
The catalytic oxidations in soils have also been found to be pro-
portional, within certain limits, to the percentage of manganese
contained therein,and to be susceptible of stimulation by the
addition of manganese compounds in much the same way as
stimulationin the activity of plant oxidases is produced by the
application of manganese salts.
Certain otherinvestigatorshave also studied the effectson the
solubilityof plant nutrientconstituentsof soils produced by the
application of manganese compounds. BERNARDINI (8), for
example, has shown that manganous chloridcauses a mobilization
of calcium and magnesiumfromboth soils and mineralsilicates to
a greaterdegree than is produced by potassium,sodium,or ammo-
nium chlorides.' From these facts the conclusionhas been drawn
that plant stimulation,resultingfrom the application of man-
ganese compounds,is probablydue to indirecteffectson the inert
bases of soils. As bearing on this phase of the question, Aso (g)
also pointed out that while a 4i . 8 per cent increasein the yield of
ricewas obtainedfromthe firstapplication of manganous sulphate,
similarapplications to the same soil the followingyear produced
only a 2. 2 per cent increase.
I See NOTTIN, Ann. Sc. Agron. Ser 4. pp. 1-12. 1913.
I914] KELLEY-FUNCTION OF MANGANESE 215

The toxic effectabove mentionedusually manifestsitselfby a

dying-backof the growingtips and the yellowingof the leaves.
LOEW and SAWA (5) hold that the activityof the oxidizingenzymes
of plants may become excessive under the influenceof sufficiently
great amounts of manganese, resultingin the auto-oxidation of
the chloroplastids,thus destroyingthe greenpigment. SALOMONE
(IO), on the other hand, found evidences of plasmolysisin wheat
that had been fertilizedwith manganesedioxide at the rate of fifty
kilograms per hectar. BRENCHLEY (ii) also observed a toxic
action frommanganous sulphate when it was applied in the very
low concentrationof one part per one hundred thousand parts of
culture solutions. When it is remembered,however, that many
plants vegetate normallyin culturesolutionsvaryingrather.widely
in ionic-concentration, without showing evidences of plasmolysis,
in otherwords,the ionic-concentration of the cell sap considerably
exceeds that of ordinary culture solutions, it seems improbable
that the addition of such small amounts of manganous sulphate
would produce plasmolysis as a direct effect.
From the foregoingbrief resume2of the experimentson this
question,it will be seen that two different sets of views have been
held by which the action of manganese on soils and plants has
been explained. These may be brieflystated as follows: (i) man-
ganese stimulates the necessary oxidations going on in soils and
plants through the activation of the oxidizing enzymes, etc.;
(2) the application of soluble manganese bringsabout plant stimu-
lation by renderingsoluble, and thereforemaking available, the
essentialplant food of soils.
Under the firstnamed of these theoriesmay be included the
stimulatedoxidations,in both soils and plant tissues, the former
probablybeing referableto the action on certainmicroorganisms,
as wellas on soil catalysis,whilethe latterhas to do withthephysio-
logical processeswithin the cell. Viewed in the light of the last
named theory,the effectsof manganese on plants are considered
to be indirectand due to its increasingthe solubilityof the mineral
constituentsof soils.
2 A more completebibliographyof literaturedealing with this subject will be

foundin a paper by the writerpublishedas Bulletin26, Hawaii ExperimentStation.

216 BOTANICAL GAZETTE [MARCH

A study of the literatureon this subject, embracingas it does

experimentswith a considerablerange of plants, reveals discrep-
ancies,and, in theopinionof thewriter,casts doubt on the adequacy
of eitherone or both of the above mentioned conceptions as fur-
nishing sufficient basis for a completeexplanationof the function
of manganesein plants. In this connection,the application of the
principlesof plant physiology,particularlyosmosis in its bearing
on the absorption of chemical substances from soils; naturally
raises the question of the influence of soluble manganese on
the selective absorption of the necessary nutrient elements.
SCHREINER and SKINNER (I2) have shown,forexample,that small
amounts of cumarin materiallyalter the absorptionof potash by
wheat seedlings; that quinone modifiesthe osmotic absorptionof
phosphoric acid; and that vanillin and dihydroxystearicacid
exerta noteworthyinfluenceupon the absorptionof nitrates. Up
to the presenttime the fundamentalsteps and the specificsof these
reactionshave not been elucidated. The exact "how" as yet can
onlybe inferred. With regardto the specificeffectson the osmotic
phenomenon produced by small amounts of inorganic elements
occurringin soils, other than the so-called plant food elements,
little indeed is known. It is reasonable to suppose, a priori,
that some osmotic effectwould be produced. In fact,it seems
improbable that the presence of any considerable amount of an
electrolytein the nutrientsolution would exert a strictlyneutral
effect.
For a numberof years the writerhas studied the effectof man-
ganese on plants. A preliminaryreportof thisworkwas published
in i908 (I3), in whichit was shownthat certainsoils of Oahu, used
forpineapples, containabnormallyhigh percentagesof manganese.
On these areas the pineapple undergoesthe phenomenonof chloro-
sis, the leaves becomingyellow and the fruitpink in color through-
out the entire period of its growth. The growthof the plant is
stuntedand the fruitproducedis inferiorin size. Furtherinvesti-
gation of this question has led to a study of the functionof man-
ganese in plants in general. the results of which, in view of the
prominenceof the question. appear to be of sufficient interestto
warranta briefdiscussionat this time.
1914] KELLEY-FUNCTION OF MANGANESE 217

observations
Experimental
The macroscopic appearance of a number of plants when
grownon manganiferoussoil is characteristic. Pineapples develop
chlorosis at an early age, fromwhich they never recover. The
effectsare firstnoticeable by a yellowingon the margins of the
leaves or the developmentof yellow spots which soon spread over
the entire leaf. The growing tips die back, the leaf margins
becoming brown. Usually such plants produce small fruitof an
abnormallypink color. Pineapple roots, instead of maintaining
theusual pointedgrowingtip,are oftenfoundto be greatlyenlarged,
sometimesto the size of a lead pencil. The roots of otherplants
also become modifiedto some extent. Those of Panicum molle
are foundto be abnormallywoody. Others are similarlyaffected.
Corn makes poor growth,the leaves turningbrown at an early
stage and the stalks taking on a deep purple color. Barley, oats,
and rice are likewisestuntedin growth. Of the Leguminosae, the
cowpea (Vigna catjang), in particular,is very sensitive to man-
ganese; the lower leaves become brown, die back fromthe tips,
and fall away. The pigeon pea (Cajanus indicus) is affectedsome-
what similarly. Onions also die back fromthe tips and produce
very small bulbs.
Some species, however,appear not to be affected. The Agave
sisalana shows no effects; sugar cane is not greatlyaffected,and
cotton and tobacco, although somewhat retarded in growth,are
not otherwiseaffected. Waltheriaamericana,the sow thistle,and
a species of Crotalaria grow as weeds on the manganiferoussoils
and are in no way hindered in their development. These illus-
trationscould be greatly enlarged, but are sufficientto indicate
that the apparent effectsof manganese in different species are far
frombeinguniform.
MICROSCOPICAL STUDIES.-From a microscopicalstudy of the
several parts of these plants, it was foundthat in those instances
where manganese exerts a toxic effect,the cell walls of the root
cortexbecome brown,and in some instancescontain minutegran-
ules of manganese dioxide. In a few instances the cells having
brown cell walls extendinto the centralcylinderof the root, and
2IS BOTANICAL GAZETTE [MARCH

it is not uncommonto finda thickened tissue lying next to the

cell walls in the angles of the cells. In the more advanced stages
of chlorosisthe protoplasmiccontentsof pineapple leaves become
contracted into formlessmasses and appear to undergo partial
decomposition. Normally, the palisade cells (14) of this species
contain not more than mere traces of chlorophyll,being essentially
water-storagetissue. Under the influenceof manganese the liquid
contentbecomes coagulated and draws away fromthe cell walls;
likewise the protoplasm in the chlorophyll-bearing cells breaks
away fromthe cell walls in places, contractinginto an irregularly
shaped mass. Plasmolysis, therefore,takes place, and in a few
instancesthe nuclei become brown.
At the beginningof the chlorosisthereis a fadingof the green
color withoutany othernoticeablechange; but in successivestages
the chloroplastidsgradually become fewerin numberand smaller
in size until finally they almost completely disappear. Simul-
taneously with the fadingof the chlorophyll,a diminutionin the
starch contentstakes place, and as the chlorosisdevelops, less and
less starch is found adheringto the chloroplastids,until finallyit
entirely disappears. In the chlorotic plants the palisade cells
also seem to be given over to the storage of calcium oxalate, and
the occurrenceof this metabolic by-productis greatly increased
in practicallyall parts of such plants.
OXIDIZING ENZYMES.-The -view that manganese exerts an
influenceon plants, throughstimulatingthe auto-oxidations (so
indispensable to living matter), gives interestto a study of the
oxidizing enzymes occurringin plants frommanganiferoussoils.
Accordingly,the oxidase and peroxidase activities of infusions
obtained by crushingthe leaves were measuredempiricallyby use
of the guiacum and aloin reactions. The resultsshowed that while
infusionsfrom some chloroticplants contain vigorous oxidizing
enzymes.frequentlysuch infusionsgive much weaker oxidase and
peroxidase reactions than normal plants. These tests have been
applied to infusionsfromall parts of pineapple plants, comingfrom
soils containingmanganese in amounts varyingfromo.oi to 9.75
per cent,and withplants at all ages and in the various stages in the
developmentof chlorosis; but when applied to a large series of
I9141 KELLEY-FUNCTION OF MANGANESE 2I9

samples no correlationwas found between the percentageof man-

ganese in thesoil or the degreeof chlorosison the one hand, and the
activityof the oxidizingenzymeson the other.
These tests have also been made with various other plants,
some of which show toxic effectsfromthe manganese, and with
still others that appear not to be affected. On the whole, the
resultsagain fail to show any relationbetween the activityof the
oxidizing enzymes and the percentage of manganese in the soil,
or the toxic effects.
As is well known,the activityof oxidizingenzymesin different
specimensof a given species of plant is by no means uniformwhen
grown on normal soils. It has been found, for instance, that
extractsobtained fromthe freshleaves of normal sugar cane and
pineapples, respectively,varied in their oxygen-carryingpower
between wide extremes. It is also known that pathological dis-
turbances,caused by attacks of Aphidae, the mosaic disease of
tobacco, etc., are also associated with accelerated oxidation. The
autumnal yellowingof plants, incidentto maturity,and the devel-
opmentof yellowspots on certainplants,have likewisebeen shown
by WOODS (I5) to be associated with an increased activity of the
oxidases and peroxidases.
It seems reasonable to suppose, therefore,that while man-
ganese has the power of increasingthe oxygen-carrying power of
oxidases,and consequentlymay, in this way, to some degreebring
about plant stimulation,the phenomenonof chlorosiscannot be
completely explained on the basis of excessive auto-oxidation.
The development of chlorosisunder the influenceof manganese
is veryprobably the resultof physiologicaldisturbancesof a more
deep-seatednature.
COMPOSITION OF THE ASH.-A number of ash analyses have
been made forthe purpose of determiningthe effectsof manganese
on the absolute, as well as the relative,absorptionof plant food
elements. The materials for analysis were selected with the
greatest care, so as to secure representativeplants of the same
age and stage of developmentin a given species. The resultsare
recorded in table I.
The data given in table I bringout some interestingfacts. In
220 BOTANICAL GAZETTE [MARCH

TABLE I
ASH ANALYSIS OF PLANTS GROWN ON NORMAL AND MANGANIFEROUS SOILS

Phosphorica
Plants analyzed Manganese Lime Magnesia Total ash
(MnOj (CaO) (MgO) acid (PA.0s)

Pineapple leaves 5 monthsold: P ct. P ct. P ct. P ct. P ct.

Manganiferoussoil. . . . . . . . . . . 2.4I 9. 0I 5.70 2.8I 9.94
Normalsoil.................. I .70 7.I4 7.60 3 . 57 7. I4
Pineappleleaves i8 monthsold:
Manganiferoussoil..... ........ 2. o8 I5.66 7.9I I.66 7.98
Normal soil.................... I .40 7.00 6.98 2.70 6.24
Pineapple stalk 5 monthsold:
Manganiferous soil ............ . 25 36.42 2.60 6. I2 8.85
Normal soil.................... 25 23.87 5.82 8.86 5. I2
Pineapple stalk 2 yearsold:
Manganiferoussoil..... ........ .80 I4.36 7 . 75 6.70 7.78
Normal soil.................... .20 I2.96 5.78 8.36 6.6o
Corn stover:
Manganiferoussoil..... ........ .40 8.6o 7.64 5.I7 7 . 55
Normal soil................... . I5 3 .45 4.60 7.56 9. I8
Cowpea vines:
Manganiferoussoil............. 3.07 22.72 6.93 2.90 I3.73
Normal soil .................... .87 I7.IO 9. I3 5.03 I3.44
Cowpeas:
Manganiferoussoil............ . I5 I .47 4.I5 I5.89 3 .93
Normal soil .................... . 1.
II3 6.9I 22.7 I 3 .53
Paspalum orbiculare:
Manganiferoussoil............ 2.20 5.30 3.46 2.54 7.88
Normal soil .................... I .05 6.70 5 .05 2.34 7. I6
Guava leaves:
Manganiferoussoil...... ....... .82 43.02 3 .73 4.47 7.66
Normalsoil.................... .28 22.98 IO.46 7.70 6.73
Guava wood:
Manganiferoussoil...... ....... I1.20 52. IO I. 5I 4 -33 5.87
Normalsoil .................... I5 26. I7 7.22 6.97 4.02
Sugar-caneleaves:
Manganiferoussoil .... . . . 45 I4.33 4.4I 3.2I 5 . 57
Normal soil.................... .6o i6.6o 6.07 4.03 4.85
Crotalaria:
Manganiferoussoil..... ........ I .40 30.80 7 .79 4. I2 6.77
Normal soil .................... .05 I9.00 2I.05 9.I3 6.28
Peanut leaves:
Manganiferoussoil............. .25 35. 24 5.6I 4-49 IO.45
Normal soil.................... .04 39.4I IO0.99 4 -97 9.95
Peanut stems:
Manganiferoussoil..... ........ 2 .56 I4.80 9.04 2.00 II.I5
Normal soil. . . . . ...... . ........ 32 2I .76 I9.02 4.8I 7. I 2
Ironwood (Casuarina equisetifolia)
needles:
Manganiferoussoil. . .66 37.50 4.57 i.62 ........
Normalsoil.................... T. 43. I2 7- 75 4.23 ........
Olive leaves:
Manganiferoussoil..... ........ .64 26.32 I .84 2.63 ........
Normalsoil.................... .86 I 7.80 I .53 2.78 ........
Waltheriaamericanaleaves:
Manganiferoussoil. ............ 8.70 3I .30 3.8I 2.80 ........
Normal soil.................... .82 29.62 5.44 3.8I ........
I9I4] KELLE -FUNCTION OF MANGANESE 22I

TABLE I-Continued

Plants analyzed Manganese Lime Magnesia Phosphoric ash

(Mn304) (CaO) (MgO) acid (PD05)Total

Waltheriaamericanastems:
Manganiferoussoil............. 56 I4.97 II .70 3.67 3. 58
Normal soil.................... 45 I3.70 II .30 3.46 3 73
Broom-cornleaves:
Manganiferous soil .2.24 I 2. o8 6.75 .94 ........
Normal soil................... .6o 5-44 3.52 I.38 ....
Broom-cornstalks:
Manganiferoussoil. I .36 2. I2 3. I2 I.02 ........
Normalsoil .................... 52 I.88 2.5I 3.72 ....
Tobacco stems:
Manganiferous soil. 57 9.47 4. o8 2. o8 7.02
Normal soil.................... T. I.88 3.27 7. 35 8. So
Pigeon-pealeaves:
Manganiferoussoil............. I .43 i6. II 3.25 4.04 I2. I7
Normal soil . ..........7....... 5 7.86 7.66 Io.85 7.84
Pigeon-peastems:
Manganiferous soil . . . . . . . . . . . . . 99 I5.29 2.82 4.25 5.8i
Normal soil .................... T. 3 .79 4.39 9.34 6.22
Oat straw:
Manganiferoussoil ............. .86 9. I5 5. i6 .73 I0.27
Normal soil .................... T. 3.40 3. I9 8.8I I I .73
Wheat straw:
Manganiferoussoil.22 .............2 4.5 I 4.03 2.96 8.9I
Normalsoil.................... . 2.09 2.74 5.56 i6.27
Mango leaves:
Manganiferous soil. . . . . . . . 2. i6 33. I2 2. I 2.89 9.24
Normalsoil ................ . .24 I7.13 4.90 5 .43 8.2I

practicallyeveryinstancetheabsorptionofmanganesewas increased
on the manganese soil. The analyses furthershow that thereis a
pronounced differencein the percentagesof lime, magnesia, and
phosphoricacid in the ash of plants fromthe two classes of soils.
Some of the plants were not visibly affected; others little so;
while still othersshowed a pronouncedtoxic effect;but uniformly
throughouttherewas a tendencytowardan increasedabsorptionof
lime on the one hand, and a decreased absorptionof magnesia and
phosphoricacid on the other.
Under certain conditions the ratio of lime to magnesia seems
to be of considerableimportanceto plants. In order to bringout
more clearly the relations between these two constituentsof the
ash, the relative amounts of lime and magnesia absorbed from
normal and manganiferoussoils, respectively,have been recal-
culated fromthepreviousash analysesand are presentedin table II.
An inspectionof the data in table II shows that in almost every
222 BOTANICAL GAZETTE [-MARCH

TABLE II
THE RATIO OF LIME TO MAGNESIA IN PLANTS (MAGNESIA CONSIDERED AS I)

From From From From

Kind of plant manga- normal Kind of plant manga normal
nf
soil niferous Soil
soil
nieosu soil
Pineapple leaves: Ironwoodneedles 8. 20 5. 56
5 monthsold. I .58 94 Olive leaves........ I4.30 II.63
i8 monthsold ..... I. 98 I . 00 Waltheria americana:
Pineapple stalk: Leaves ... 2I 5.44
5 monthsold ..... I4.00 4.10 Stems........... I. 28 I. 2I
i8 monthsold ..... I. 88 2. 24 Broom corn:
Cornstover......... .12 .75 Leaves. .......... I 99 1
.I54
Cowpeas: Stems. 67 .74
Vines............. 3. 28 i. 86 Tobacco stems...... ..3 2 32 57
Seed ............. 35 . I6 Pigeon peas:
Paspalum orbiculare.. I . 53 I*32 Leaves ........... 4.69 I.02
Guava: Stems.....5.. . .42 .86
Leaves . .......... II.53 2.19 Oat straw. ......... I*97 I.07
Stems. 34.50 3.62 Wheat straw. ....... I.I2 .76
Sugar cane leaves .... 3.25 2.73 Mango leaves ....... It.40 3 49
Crotalaria. .......... 3 95 .90
Peanut:
Leaves ............ 6.28 3.58
Stems ............ I.63 I.14

instance the ratio of the absorbed lime to absorbed magnesia was

increasedunder the influenceof manganese.

Discussion
In this investigationit has been shown that differentplants,
whengrownon manganiferoussoils,are affecteddifferently.Some
species are stunted in growthand die back from the tips of the
leaves, which turn yellow or brown,and sometimesfall off,and a
general unhealthy appearance results. Other species appear to
be unaffected,and, so far as can be judged, vegetate normallyin
the presence of manganese. Microscopicinvestigationsshow that
in certain instances the protoplasm undergoes changes. Occa-
sionallyit draws away fromthe cell walls and the nuclei become
brown. There is a manifestchange in the protoplasmiccontents
of the roots.
The chlorophyllin a numberof plants is affected;in pineapples
it undergoesdecomposition. Simultaneouswith the destructionof
chlorophyll,starch formationceases.
The activityof the oxidizingenzymesin plants is hereinshown
to. bear no relation to the destructionof chlorophyllunder the
191I4] KELLEY-FUNCTION OF MANGANESE 2 23

influenceof excessive manganese. While the oxidases generally

contain manganese as a normal constituent,or at least manganese
is closely associated with the oxidases, and at the same time their
oxygen-carrying power is accelerated by the presence of man-
ganese salts, the foregoinginvestigationsshow that there is no
correlationbetween the phenomenon of chlorosis in pineapples
and the activity of the oxidizing enzymes. The decomposition
of chlorophyllin this case, therefore,is not due to excessive auto-
oxidation. This does not imply, however,that accelerated auto-
oxidationin plants is withouteffect.
From the ash analyses it was found that manganese was
absorbed in considerablequantities, and in nearly every instance
was greaterin the plants frommanganiferoussoil. These analyses
also show that a disturbancein the mineral balance takes place.
The percentageof lime is increased,while the percentageof mag-
nesia and phosphoricacid is decreased. Some of the plants ana-
lyzed showed a markedly toxic effectfrom the manganese,while
othersappeared to be unaffected;but in practicallyeveryinstance
a modificationof the mineral balance was observed, and this was
found to follow the same directionin all species. The ratio of
absorbed lime to the absorbed magnesia was increased under the
influenceof manganese, regardlessof whetherthe plant showed a
toxic effector not:
It is claimed by LOEW (i6) and othersthat various plants are
affecteddifferently by differentratios of lime to magnesia; certain
species of plants vegetate most advantageously when the ratio is
one to one; whereas still other species grow best when the ratio
is one to three,etc. In the publicationsdealing with this subject,
however,mentionis usually made of the effectsproducedupon the
appearance of plants, while but few ash analyses have been made
in this connection. From the data at hand, however,it appears
that where the physiologicalbalance between lime and magnesia
is disturbed,a correspondinginfluenceis broughtabout in the com-
positionof the ash.
Under the influenceof manganese,plants automaticallymodify
themselvesin regardto the absorptionof these two elements,and,
as is believed, it is not so much the absolute amountof calciumand
224 BOTANICAL GAZETTE [MARCH

magnesiumin a given soil as the ratio betweenthesetwo substances

in solution in the soil moisturethat determinesthe physiological
effects. In the case of plants grown on manganiferoussoil, the
relative amounts of lime and magnesia actually absorbed become
greatlydifferent fromthose absorbed fromnormal soils, regardless
of the amounts of these elementspresentin the soil.
From these evidences,we may believe, then,that an important
effectofmanganeseis of an indirectnature,beingdue to its bringing
about a modificationin theosmoticabsorptionoflimeand magnesia,
and that the toxic effectsare chieflybroughtabout by this modifi-
cation, ratherthan as a direct effectof the manganese itself. As
has been mentionedalready, not all species of plants are equally
sensitive to modificationsin the lime-magnesiaratio; and from
some recentexperimentsof GILE (I7) it seems that the effectsof a
modificationin this ratio are, to some extent,dependentupon the
concentrationof the culture solution. Likewise, differentratios
are best suited to different species. Therefore,th&effectof man-
ganese may be very differenton different soils and with different
species of plants. With certain plants it is toxic on certain soils
forthe reason that the absorbed lime and magnesia are thrownout
of theiroptimumratio forthis plant, while in othersit may exert
a stimulatingeffectby bringing this ratio more nearly to its
optimum.
The small amounts of manganese in natural soils, therefore,
probably performa twofoldfunctionin plant growth: (i) it acts
catalytically,increasing the oxidations in the soil and acceler-
ating the auto-oxidationsin plants; and (2) it tends to modifythe
absorption of lime and magnesia, perhaps by partially replacing
calcium from insoluble combinations,but especially, througha
direct effecton the osmotic absorption of lime and magnesia,
increasingthe formerand decreasingthe latter.
The absorption of phosphoricacid is likewisedecreased in the
presence of manganese. By referenceto the preceding table of
ash analysesit will be seen that frequentlythe ash of a given species
of plant frommanganese soil was found to contain not more than
one-halfas much phosphoricacid as fromnormalsoil. The inter-
ferencewith the absorptionof phosphoricacid would also tend to
I9I4] KELLEY-FUNCTION OF MANGANESE 225

bringabout stuntedgrowthand mightbe sufficient to account for

the differencein size of the plants from the two classes of soil.
Studies on the solubilityof these soils have shown the manganese
to be slightlysoluble in water, but markedlyso in dilute organic
acids. Phosphoric acid coming into solution in the soil moisture
would tend to be precipitated by the manganese as manganese
phosphate, a compound, which can be dissolved only with diffi-
culty,and thereforethe absorptionof phosphoricacid would thus
be hindered.
Referencehas already been made to the factthat certainorganic
substances exert appreciable influenceon the rates of absorption
of certain of the necessary elements. Here it is shown that a
similaraction is to be ascribed to manganese. How is this fact to
be explained?
During recent years the protective action of salts shown to
obtain in animal experimentsby LOEB (i8) has been found to
apply also to plants.
OSTERHOUT (i9) has shown fromwater cultures,for instance,
that some of the essential elementsmay be toxic unless properly
balanced by definite concentrationsof certain other necessary
elements. In otherwords,some at least of the essentialinorganic
elements seem to play both a nutritive and a protective role.
Quite recently OSTERHOUT (20) has been able to throwconsiderable
lighton -themechanics of the protectiveaction. By the use of an
ingeniousdevice he determinedthe electricalconductivityof living
protoplasm, both before and after it was placed in solutions of
sodium and calcium chlorides, etc. The results show that the
electrical resistance of the protoplasm becomes greatly reduced
upon standing in sodium chloride solution, thus indicating that
sodiumhad been absorbed. But when the protoplasmwas placed
in a solutioncontainingsodium and calcium chlorides,of the same
ionic concentration,no such loweringof the resistancetook place.
Calcium hinders,then, the passage of sodium throughliving pro-
toplasm; and thereforemust be looked upon as loweringthe per-
meability. OSTERHOUT has furthershown this phenomenon to
be reversible,and thereforethe presenceof one salt may actually
cause an increasein thepermeabilityto others. In certaininstances
226 BOTANICAL GAZETTE [MARCH

he found the protoplasm to undergo visible changes, which,how-

ever, were not necessarilyinjurious to it. Protoplasm,being of a
colloidal nature, then, becomes physico-chemicallyaltered when
placed in certain solutions; and such alterations,in turn,affect
its permeability.
By the application of the above named conceptions,we may
inferthat when plants are grown on manganese soils, the soluble
manganese, coming into contact with the root hairs, is at first
absorbed, up to a certain point, formingcombinationswith the
protoplasm. Once these combinations are established, the per-
meabilityof the protoplasmbecomes altered,wherebythe absorp-
tion of lime is facilitated,while that of magnesiumis hindered.
Manganese, then, may be looked upon as forminga combination
with the protoplasm which materiallyalters the relative absorp-
tion of lime and magnesia. Not all plants are equally affectedby
a variation fromthe normal ratio of lime to magnesia, and there-
foreunder fieldconditionstoxic effectsmay be produced in some
plants, stimulationin others,while with still othersthe effectsmay
be neutral.
The experimentalfacts presentedin this paper and the inter-
pretationsmade in no way negative the work previouslydone on
this subject. The failureto observe a more active catalytic oxi-
dizing reaction in plants fromhighlymanganiferoussoil does not
argue that manganese is incapable of stimulatingthe activity of
oxidizingenzymes. The normal soils of Oahu, fromwhich plants
were taken for comparison,contain small amounts of manganese,
which,moreover,was foundto be absorbed to a considerableextent
by all theplants studied. It is probable,therefore,that theabsorp-
tion of manganese fromthe normalsoils of the Islands is sufficient
to produce maximum stimulationin oxidase activity.
HAWAII EXPERIMENT STATION
HONOLULU

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