Food Sec.

(2011) 3:93105
DOI 10.1007/s12571-011-0112-9

ORIGINAL PAPER

Stress-adapted extremophiles provide energy

without interference with food production
Ray A. Bressan & Muppala P. Reddy & Suk Ho Chung &
Dae Jin Yun & Lowell S. Hardin & Hans J. Bohnert

Received: 10 November 2010 / Accepted: 30 January 2011 / Published online: 19 February 2011
# Springer Science+Business Media B.V. & International Society for Plant Pathology 2011

Abstract How to wean humanity off the use of fossil fuels good sense. It is clear that not every advertized energy source
continues to receive much attention but how to replace these can lead to a sustainable, humane and environment-friendly
fuels with renewable sources of energy has become a path out of a future energy crisis. Our proposal is based on two
contentious field of debate as well as research, which often assertions: that the use of food crops for biofuels is immoral,
reflects economic and political factors rather than scientific and that for this purpose using land suitable for growing crops
productively is to be avoided. We advocate a focus on new
extremophile crops. These would either be wild species
R. A. Bressan adapted to extreme environments which express genes,
Department of Horticulture and Landscape Architecture,
Purdue University,
developmental processes and metabolic pathways that distin-
W. Lafayette, IN 47907-1165, USA guish them from traditional crops or existing crops genetically
modified to withstand extreme environments. Such extrem-
R. A. Bressan : M. P. Reddy ophile energy crops (EECs), will be less susceptible to
Plant Stress Genomics Research Center,
stresses in a changing global environment and provide higher
King Abdullah University for Science and Technology,
Thuwal 23955-6900, Saudi Arabia yields than existing crops. Moreover, they will grow on land
that has never been valuable for agriculture or is no longer so,
S. H. Chung owing to centuries or millennia of imprudent exploitation.
Clean Combustion Research Center,
King Abdullah University for Science and Technology,
Such a policy will contribute to striking a balance between
Thuwal 23955-6900, Saudi Arabia ecosystem protection and human resource management.
Beyond that, rather than bulk liquid fuel generation, combus-
D. J. Yun : H. J. Bohnert tion of various biomass sources including extremophiles for
Department of Plant Biology and Department of Crop Sciences,
University of Illinois at Urbana-Champaign,
generating electrical energy, and photovoltaics-based capture
Urbana, IL 61801, USA of solar energy, are superbly suitable candidates for powering
the world in the future. Generating electricity and efficient
R. A. Bressan (*) : H. J. Bohnert (*) storage capacity is quite possibly the only way for a
Division of Applied Life Sciences, WCU Program,
sustainable post-fossil and, indeed, post-biofuel fuel economy.
Gyeongsang National University,
Jinju 660-701, South Korea
e-mail: bressan@purdue.edu Keywords Alternative crops . Bioenergy generation .
Extremophiles . Abiotic stress tolerance . Food or fuel
R. A. Bressan
e-mail: ray.bressan@kaust.edu.sa

H. J. Bohnert Introduction
e-mail: hbohnert@illinois.edu
The current massive use and dependence on fossil fuels will
L. S. Hardin
Department of Agricultural Economics, Purdue University, eventually lead to their depletion but, more importantly, the
W. Lafayette, IN 47907-1165, USA present applications and uses of fossil energy sources
94
contribute significantly to altering the environment.
Resource exploitation is accelerating as industrialization
reaches all regions of the earth, and as people world-wide
work to improve their lives. Industrialization, population
increase and the lack of enforceable standards in most
countries has also led to widespread abuse of the land and
environmental degradation. These outcomes are consequen-
ces of a near single-minded focus on profit, accompanied
by significant income inequality. Manipulations and subsi-
dies have hidden the true cost of energy generation models
that have emerged over the last 200 years. Dependency on
fossil fuels, leading to environmental degradation and global
changes, seems to go ahead largely unchecked as developing
countries achieve parity with the more developed countries.
Continuing along this path could lead to a global crisis that,
without arbitration, has the potential to result in momentous
cultural, social and economic tensions.
Growing awareness of the facts and certain consequen-
ces of continuing along the same unwise trajectory has
resulted in a number of proposals for possible remedies
about how the worlds craving for energy might be
satisfied. Among the options, the production of alternative
fuels from biomass has often been put forward as one
important component for resolving the energy crisis. Other
avenues point to harnessing wind and wave-power,
geothermal or photovoltaic energy capture as possible
remedies. All options are being pursued at present; the
final outcome will most likely be a mix of different
approaches. With a still growing human population,
judgment about whether we will achieve global energy
autarky without damaging the biosphere is not yet possible.
Being renewable in principle, biomass indeed assumes a
prominent place among the options that must receive
serious consideration. So far, however, this path for
producing bio-fuels has been sidetracked by players with
an economic interest on using food crops for ethanol
production. There are considerable flaws in the approach
and serious detrimental consequences will be the outcome
should this path be followed, unmodified.
Assigning a significant proportion of the North American
corn crop to ethanol production in 2007, coupled with
environment-based failure of rice and other critical crops,
and increased demand for high caloric foods in rapidly
developing countries drastically increased food prices
(Mitchell 2008). Soaring food prices were accompanied
by social unrest leading, in isolated cases, to riots in
several countries (Brown 2008). This has generated what
might be termed a dress rehearsal for the possible
consequences of the precarious future of world food
security. Two trends are driving humanity to a seemingly
unavoidable food crisis that is virtually not acknowledged
in its impact and severity. Population increase continues at
a pace that will add some 2.5 billion people by the year
R.A. Bressan et al.
2050 to the 6.5 billion already on the planet. Second and
compounding the problem is that people in developing
countries are successfully crossing into a middle class
status, accompanied by increases in per capita caloric use.
Information technology also has provided virtually every-
one with the means to appreciate the wealth that had
previously been reserved to a few and to now seek
participation. This almost invisible crisis is greatly over-
shadowed by the inevitable energy crisis which is
demanding solutions, even when the detection and
exploration of new fossil fuel sources will postpone the
unavoidable eventual decline. That it even could be
considered to use food to produce energy in a world
where almost 1 billion people are undernourished speaks
of insane, inconsiderate objectives. The lessons of 2007
should be heeded, viz, that increased energy supply cannot
rely on food biomass (Patzek 2007).
The problem of the first generation ethanol biofuels from
corn can be readily assessed from the statistics in energy
consumption [Key World Energy Statistics, 2009, Interna-
tional Energy Agency]. The energy consumption per capita
per year in developed countries, e.g., USA is 7.75 ton of oil
equivalent (TOE), which corresponds to 212,330 Kcal/ca/day
based on 2007 energy consumption statistics. Even mid stage
developing countries such as Korea, Taiwan, and Saudi
Arabia consume 125,700, 131,780, and 170,140 Kcal/ca/
day, respectively. Average global everyday food consumption
per person is about 2,400 Kcal/ca/day. This demonstrates that
we are consuming 50100 times our capacity for total present
food energy production as the sum of fuel energy sources.
Considering the portion of corn in our foods, even accounting
for the fact that corn is mainly utilized for feeding animals for
meat production, the first generation biofuel production from
corn or, indeed, all food, is not a viable option for solving the
energy predicament.
We shall discuss the following considerations about the
generation and use of biomass as components of the
resolution of this crisis: (1) Non-edible biomass must be
the source of any large scale biofuel production that enters
the global supply system (World Development Report
2010). (2) Liquid or alcoholic fuels from bioconversion
mechanisms can play important roles in areas not connected
to supply grids, where small scale production systems can
be employed to provide energy. However, for advocating
entry into a global industry from cellulosics there are
sufficient data suggesting that liquid fuels from renewable
sources cannot be produced in an energy-dense and
environmentally tolerable way (Patzek 2007). Certainly,
liquid fuels will have a future for specific purposes, such as
kerosene used in airplanes, and production systems based
on algal biomass are promising. (3) Combustion rather than
chemical or biochemical conversion to liquid fuels is the
more likely successful route to energy production. The gain
Stress-adapted extremophiles provide energy without interference
of energy in such a way is several-fold more efficient than
ethanolic fermentation of starch- or cellulose-based fuels
(McKendry 2002; Campbell et al. 2009). (4) Finally, we
advocate the generation of new crops, which can be based
on established crops but which can also be found among
wild species (Bressan et al. 2001; Gutierrez and Ponti 2009;
Weyens et al. 2009). Generation of these crops should use
one of two strategies: transgenic means or breeding
programs that are based on genome information. Especially,
we consider as one important answer for improving the
generation of biomass the engineering or enhancement of
genes, processes and metabolic pathways that are used by
species that are naturally adapted to thrive in extreme
environments (Office of International Affairs 1990; Bressan
et al. 2001; Amtmann et al. 2005).
Non-food biomass
In principle, there are two major sources of non-edible
biomass. One is made up of the non-edible portions of
major crop plants, e.g. wheat and rice straw or corn
stover (Graham et al. 2007). This practice may, over
time, lead to soils that become depleted in carbon. The
second non-food source of biomass, which is substantial
and most likely to provide an economically advanta-
geous route, are specialty crops with large biomass
yields in environments less suitable for extant crops.
Whereas we foremost consider land areas here, generat-
ing biomass or useful chemical compounds, such as
lipids, from aquatic systems is another option. Harness-
ing the productivity of the sea may be of importance in
the future. For example, unicellular algae in ponds, or
ocean-based macroalgal farms, have been proposed as
viable biomass/energy sources and pharmaceutical pro-
duction areas (Beer et al. 2009).
An important resource is also the biomass that may
conceivably be produced from the vast areas of land that is
agriculturally of low quality because it is characterized by
stressful environments to which our crops have never been
adapted through breeding. When we point to the potential
productivity of marginal lands, we are not picturing the
extension of agriculture to untouched valuable ecosystems
but to areas that have been degraded by agriculture in the
past. Areas in this category can be found in sub-tropical and
tropical climate zones across the globe, in the African Sahel
zone, central Asia, and indeed also in the Americas north
and south. Such degraded, low productivity land is
increasing world-wide. In this sense we are advocating the
recovery of degraded ecosystems using species that are
naturally capable of succeeding in such environments.
There are many regions world-wide where extremophiles
may be grown: in a general sense these would be areas
95
where subsistence farming is practiced with minimal gains
and high frequency of disastrous harvests that require
international support efforts. A world with nine-billion
humans will have to organize the globalization of how to
grow energy crops while providing food to those who grow
these crops. While irrigation is not strictly excluded, we
envision non irrigation as the environmentally prudent
production scheme that optimizes greening of degraded
lands in a sustainable way.
This potentially useful biomass has received little
attention. Only recently, attempts have been initiated by
organizations such as CIMMYT to generate from high-
yielding lines of maize the progeny that may be useful in
drought-prone areas of Africa (Messmer et al. 2009). Rather
than adapting existing crops, generating alternative crops
that could be used strictly for biomass production on
marginal soils in stressful environments has been proposed
before with, for various reasons, little resonance (e.g.,
Odum 1974; OLeary 1984; Glenn and OLeary 1985;
Mizrahi and Pasternak 1985; Hendricks and Bushnell
2008). It is time to reconsider. Table 1 lists species growing
well under highly stressful conditions and includes data on
biomass or seed/fruit yields, although in many cases the
depth of the information is not sufficient. Conditions, for
example salinity or aridity, under which these species were
grown were often not recorded in sufficient detail. Unlike
the well known fresh water crop species from tropic or
temperate zones these species have not or have only
marginally been improved by selection or breeding. Given
the plant breeders toolbox, now fortified by genomics,
such species should be targeted for improvement to convert
them into genuine new crops.
Non-food crops will gain particular importance as the
yield of crops for human consumption must be increased,
which will probably be achieved by breeding, adaptation
and growth of traditional crops on less productive land.
Also, the diversion of corn starch to biofuel production will
lead to price increases as food or fuel destinations compete.
This will generate incentives to grow food crops on
generally unsuitable, marginal land that promises at least
some return for farmers, although the potential for
increased destruction to environments exists. Such areas
represent prime regions for non-food crops, in particular for
species, as we shall outline below, that are adapted to
tolerate abiotic stresses in areas that are often nutrient-poor
as well as because scarce coverage by plants has not
enriched the soil.
Liquid fuels for transportation and energy
We find it remarkable that liquid biofuels should be
promoted as the most desirable portable fuels by present
96 R.A. Bressan et al.

Table 1 Extremophile species

bioenergy production potential Species Yield References

Biomass species mt .ha1 year1

1 Acacia spp. 13.0014.00 Singh and Toky 1995
2 Agave spp. 38.0045.00 Nobel 1991a
3 Albizia lebbek 5.3020.20 Kawahara et al. 1981
4 Arundo donax 45.0051.00 Williams et al. 2008
5 Atriplex spp. 12.6020.90 Aronson et al. 1988
6 Batis spp. 13.6017.90 Glenn and OLeary 1985
7 Beta spp. 42.0055.60 Kaffka and Hills 1992
8 Casuarina spp. 17.42 Goel and Behl 2005
9 Distichlis spp. 30.0040.00 Dakheel et al. 2008
10 Eucalyptus spp. 21.0029.00 Singh and Toky 1995
11 Juncus spp. 0.9644.00 Cleark and Jacoby 1994
12 Leucaena spp. 25.0033.00 Singh and Toky 1995
13 Miscanthus spp 18.0048.00 Petyr et al. 2008
14 Opuntia spp. 31.0041.00 Nobel 1991b
15 Panicum virgatum 11.0022 .00 Lemus et al. 2008
16 Pennisetum spp. 35.0045.00 Woodard and Sollenberger 2008
17 Populus spp. 2.8011.40 Laureysens et al. 2004
18 Prosopis spp. 4.0039.00 El Fadl 1997
19 Salicornia spp. 12.7024.60 Glenn et al. 1991
20 Sarcocornia spp. 2.0247.76 Palomo and Niell 2009
21 Sesbania spp. 8.0021.00 Evans and Rotar 1987
22 Spartina spp. 40.00 Odum 1974
23 Sporobolus spp. 30.0040.00 Dakheel et al. 2008
24 Tamarix spp. 5.0020.00 Biosaline Biomass 2004
Seed/oil yield by species Mg .ha1year1 (oil, seed* or fruit)
25 Azadiracta indica 2.67 Azam et al. 2005
26 Balanites spp. 1.60 Deshmukh and Bhuyar 2009
27 Calyophyllum spp. 4.68 Azam et al. 2005
28 Citrullus spp. 1.502.10*(seed) Yaniv et al. 1999
29 Elaeis guineensis. 4.005.00 Yusoff and Hansen 2007
30 Jatropha spp. 1.507.80*(seed) Jongschaap et al. 2007
31 Kosteletzkya spp 1.46 * (seed) Gallagher 1985
32 Moringa spp. 3.50* (seed) Morton 1991
33 Phoenix spp. 12.00 *(fruit) Samarawira 1983
34 Pongamia spp. 0.222.25 Karmee and Chadha 2005
35 Salvadora spp. 3.500*(seed) NEDFCL 2002
36 Simarouba glauca 0.901.20 Dash et al. 2008
37 Simmondsia chinensis 1.122.25 Duke 1983

research strategies (National Academy of Sciences 2008).
An examination of the energy conversion efficiencies casts
serious doubt on the benefits claimed (McKendry 2002).
Comparing the combustion of biomass in power plants with
the production of ethanol or other liquid energy reveals a
strikingly greater efficiency in favor of combustion in
electric power generation (McKendry 2002; Patzek 2007;
Campbell et al. 2009). Stored electric power, for example
where depleted batteries may be exchanged for charged
ones at gas stations, is as portable as the advertised
alcohols in almost all transportation uses.
We point out the advantage of combustion now in the
context of allowing much greater advantage to the use of
stressful environments for biofuel production. In addition,
biomass waste accumulating in production agriculture
scenarios, for example the fraction of corn stover that is not
better returned to the soil, or various cellulosic materials
and even household wastes, may be utilized with no or little
Stress-adapted extremophiles provide energy without interference
new infrastructure development. It is to be noted that the
technology of combustion of biomass through which
electricity can be generated is relatively mature, based on
the current technologies of waste incineration and coal
combustion. In such cases, the significant energy loss
during the process of the conversion of biomass to biofuel
can be minimized. The strategy will have the add-on benefit
of being applicable at different scales of power generating
units. Scaling can also be achieved by using different
energy production platforms that utilize the biomass, with
small-scale, local fermentation and biogas production as an
option alongside combustion or solar energy collection in
rural areas that are not, or not yet, connected to country-
wide electricity grids. Our arguments here include the
approximately one half of crop biomass that is not annually
converted into fiber, fruit or seed, and may lead to increased
productivity and yield when more is returned to the soil.
Novel crops can be found among existing species that
can provide biomass on marginal land in stressful con-
ditions where our favorite crops will not be productive or
die. These species offer several opportunities. First, it is not
necessary that we change these specialists in the evolution-
ary race to colonize resource-poor niches into new energy
crops. In the combustion model, they are highly suitable
biomass sources already. By using them for energy we also
can avoid the difficult and time consuming genetic
manipulations needed that would convert them into new
agronomically and culturally accepted food or specialized
liquid fuel crops. Beyond plant research, what is needed is
relatively modest infrastructure modification and the
generation of incentives to allow the market to utilize these
EECs. The largest cost, once the infrastructure is in place,
both economically and energetically will be for collecting
and delivering the non-edible biomass to combustion
centers connected to the electrical grid. Once biomass
generates electricity, the cost related to the distribution of
electricity to consumers can be much smaller as compared
to the cost associated with the distribution of biofuels to
consumers.
A second option, which would establish energy crop
farms on marginal land with naturally adapted species, can
be viewed as an extension of the first, albeit on a longer
timescale. Selecting and breeding from among the many
species that could be used has in the past been viewed as
wasteful, as agriculturists have opined that the useful
species have already been domesticated. The ease with
which we can now obtain genetic information about wild
species, and then begin to select for traits that in many
cases have already been defined at the gene level, seems to
argue for a change. Conventional wisdom about breeding is
in need of a revision.
The third option, most important for bioenergy crop
utilization in a more distant future, will be the design of
97
transgenic crops. By this, we suggest that species with high
biomass potential be genetically modified to become
adapted to stressful environments without losing their
vigor. A requirement for the transfer of multiple genes
with precisely structured control regions for tissue- and
condition-specific expression and transcript and protein
stability is within present technological capabilities. The
implementation of such projects must, however, be
removed from the short-time focus of companies and the
whims of agencies that operate in funding cycles, and be
structured more like the engineering programs of NASA.
Extremophile new crops
Most crops originated from sub-tropical wild species that
have been genetically modified over more than 8,000 years
by selection for increased yield under favorable environ-
mental conditions. Inadvertently, the emphasis on yield has
also generated lines with moderate stress adaptation
capacity in particular environments but none of our major
crops can tolerate extreme environmental stresses. Here we
will briefly discuss plants that have evolved to colonize
habitats characterized by the extreme environmental con-
ditions of water scarcity, high and fluctuating soil salinity
and extreme temperatures (see Table 1). EEC species with
high productivity in stressful environments exist and
provide vast potential for further improvement because
they have not been used in breeding and the genes
underlying their extremophile lifestyles have not been
exploited by genetic engineering. They can be productive
even on marginal soils, with low fertilizer input and little
management intervention. Sustainability and preservation
of soil and water resources is an intrinsic advantage of
such new bioenergy crops. Species that carry these
characteristics in their genetic makeup exist in essentially
all plant families. However, previous attempts at propa-
gating species in this category have focused on generating
harvestable seeds or fruits (OLeary 1984; Glenn and
OLeary 1985; Glenn et al. 1991; USDA 2002). In contrast,
extremophiles have not previously been selected for
enhanced total biomass.
Appropriately modified and managed, extremophile
plant species that are naturally adapted to grow on marginal
land under stress conditions can be exploited. The
combustible energy obtainable from extremophile species,
which additionally would stabilize soils and mitigate
climate degradation, is comparable to that for crop plants
on a weight basis. Some wild grasses, some halophytic
species and xerophytic agaves, for example, generate
biomass comparable to the dry biomass of corn stover
(Bogdan 1977; Glenn et al. 1991; Nobel 1991a; Spatari et
al. 2005).
98
The major problem of irrigated agriculturesalinity
A third problem arises in irrigated agriculture. Irriga-
tion is practiced not only in arid and semi-arid lands
but isdepending on water availability and costalso
used to increase yields in general. All too often
irrigation relies on non-renewable (fossil) underground
water supplies. Prolonged irrigation and intense agri-
culture use lead to increases in soil salinity. This is an
inevitable consequence of adding water at the surface
while there is no connection to the groundwater table.
Erosion, desertification and loss of soil follow. The
worlds approximately 160 million hectares of irrigated
land, representing about 15% of the farmed area on
earth, produce approximately one third of all food. One
third of that is affected by increasing salinity, requiring
costly intervention to reverse the damage or at least
slow down its progression (Pitman and Laeuchli 2002).
Continued use of irrigated farming in places like the
southwestern U.S. is practiced because financial incen-
tives exist in the form of subsidized, artificially low cost
of water and energy, but rising costs have already led to
areas of cropland being abandoned (Global Climate and
Energy Project 2010). Arid lands cannot sustain con-
ventional agriculture without jeopardizing food supply
and climate.
Currently, the water used for irrigated agriculture is
about 3000 km3/year. It is estimated that 1000 km3/year
would be required to produce enough biofuel through
irrigated agriculture to cover 10% of current world
energy consumption. Clearly, biofuel production
through irrigated agriculture does not present itself as a
viable option (Falkenmark et al. 2009; Kinzelbach 2009.
It appears to us that the only long-term sustainable
solution is to radically change our current agricultural
practices and grow crops adapted to extreme and
degraded environments. In a more general sense, a large
component of energy and food security will require the
use of stress tolerant EECs. These species can become
new crop species not requiring irrigation, able to grow in
degraded soils, such as those with high salinity,
tolerating temperature extremes, and thriving under
resource-poor edaphic conditions with low input
requirements.
These will be the future bioenergy crops; chosen for
their adaptation to inhabit niches that have not been
previously in the focus of agriculture. Their modification
and management for highly productive growth on land that
traditional crops cannot use will avoid competition and
support food security. Many stress-adapted species show
high photosynthetic conversion efficiency during parts of
the growing season, and have biochemical adaptations that
increase water use efficiency (WUE) without compromising
R.A. Bressan et al.
growth. In many regions that are typically viewed as arid,
sodic, or nutrient poor, plant species exist that are able to
produce surprisingly high biomass. Altogether these
regions equal or slightly exceed the total area of the worlds
cultivated regions. To provide some yardstick, regions with
high productivity numbers have been recorded for temper-
ate and tropical aquaticincluding moderately brackish
swamps at ~40 metric tons/ha per year (mt/ha x a) with
cultivated annual plants at 1025 mt/ha x a. This comparison
can be applied to the productivity of the most productive
tropical perennial plants and a few crops [e.g. sweet potato] at
5080 mt/ha x a, whereas tropical annuals attain 2535 mt/ha x
a (Westlake 2008).
Naturally stress-tolerant plantsthe untapped biofuel
resource
True extremophiles are species that grow on inhospitable,
marginal lands where they have to cope with abiotic
stresses such as drought, temperature extremes or salinity
and often combinations of these factors (Bressan et al.
2001; Amtmann et al. 2005). Such plants are also expert in
withstanding high irradiation under abiotic stress conditions
that preclude growth. Extremophiles are efficient scav-
engers for nutrients from soils that are degraded or poor
(Kant et al. 2008; Vincour & Altman 2005; Wong et al.
2006) and should require minimal inputs thus minimizing
environmental collateral damage, an often cited drawback
of the green revolution (Pimentel et al. 1973; Huang et al.
2002).
Extremophile species typically grow slowly, a strategy
that provides survival while hedging bets for reproductive
success. This yield penalty for growth under stress
conditions is often cited as the major obstacle for attempts
to grow desert-adapted species. However, the slow growth
phenotype is not a constraining genetic character because
many extremophiles are able to exhibit rapid and surprisingly
substantial growth spurts as conditions improve from deadly
for normal crops to merely stressful. Examples of such
behavior are legions; some desert plants, for example, achieve
biomass productivity under well-watered conditions that
approach those of currently studied biomass crops (Nobel
1991a). A recent documentation of growth habits of
extremophiles compared salt tolerance and growth in the
absence and presence of salt stress in 11 relatives of
Arabidopsis (Orsini et al. 2010). Some extremophiles among
these species, exhibited more rapid growth than Arabidopsis
in both stressed and control conditions.
As one example, Nobel (1991a) explored productivity
of agaves and cacti grown under cultivation. They
observed aboveground biomass productivity of 38 t.ha1
year1 under non-irrigated and 45 t.ha1 year1 with
Stress-adapted extremophiles provide energy without interference
irrigation for agave species, with the corresponding figures for
Opuntia sp. being 31 and 47 t. ha1 year1, respectively.
These values approximate to 30% of the dry biomass that is
generated by Miscanthus under well-watered largely stress-
free growth conditions (Heaton et al. 2008). Likewise,
Salicornia (glasswort), Suaeda (seablite), and Atriplex
(saltbush) can provide yields similar to alfalfa grown with
freshwater irrigation. However, comparisons of productivity
between monocultures and multispecies communities and
their contribution to future global energy supply have
resulted in widely differing conclusions (Garnier et al.
1997; Berndes et al. 2003). Salicornia species, for example,
are to some degree already used as crops (Fig. 1) even in the
absence of major breeding efforts.
These and many other examples paraphrase the funda-
mental distinction between growth, growth potential, and
the ability to sustain life under stress. Whether by selection,
genetic engineering or adapting appropriate changes to
agricultural practices, the number of extremophile species
already known can provide sufficient genetic material that
can be harnessed for biomass production, and some
extremophile species are already suitable for cultivation.
Identification of mechanisms employed by various
extremophiles opens the prospect of using molecular
genetics to merge their tolerance characteristics with
engineered or breeding-dependent increases in low-input
biomass production.
A number of biomass crops, potential EECs, are
presently promoted as biofuel raw materials, such as
Poplar (Populus sp.; Laureysens et al. 2004), Miscanthus
(Miscanthus giganteus, Petyr et al. 2008; Heaton et al.
2008), switchgrass (Panicum virgatum, Lemus et al. 2008)
or prickly pear (Opuntia sp., Nobel 1991b) and agaves
(Agave americana, Nobel 1991a) and, in the tropics,
Fig. 1 Salicornia brachiata
cultivation. Plants are grown on
saline soils irrigated with
seawater (CSMCR, Bhavnagar,
India). The insert shows fruiting
plants
99
elephant grass (Pennisetum purpureum, Woodard and
Sollenberger 2008). To some degree it has been shown that
these species can be reasonably productive on land that is too
poor for agronomic profitablility or even suitability for
subsistence farming, forcing farmers to migrate, typically
joining the urban poor. In addition, some species such as
palms (Elaeis guineensis, Lam et al. 2009) and physic nut
(Jatropha curcas, Achten et al. 2007) are productive on less
fertile land producing oil suitable for conversion into
biodiesel (Fig. 2). Oil palm monocultures in southeast Asia
are profitable and have improved living conditions, although
the cultivation is already seriously encroaching on both
natural ecosystems and land that was previoulsy used for
growing food crops. Such practices are not sustainable in the
long term.
The yield penalty conundrum is also suspect for another
reason because yield or greater EEC biomass is genetically
controlled and can be engineered. Many attempts have, in
the past, been directed towards engineering photosynthesis,
viewed as the most basic determinant of productivity.
There have been only few, if any, yield improvements by
this route. More promising, it seems, will be that we focus
on the structure of gene systems in plants with different
lifestyles and, especially, the interaction of genes with the
environment. Here we point to new data and insights that
indicate how environmental factors can alter gene expres-
sion (Jablonka and Raz 2007). Recently a number of
studies have shown that growth can be altered by
controlling hormone synthesis, distribution and, especially,
the ratios among plant hormones (Bgre et al. 2008). In
essence, the perception by a plant of what is a stressful
condition and what constitutes tolerable stress that may be
altered without incurring a yield penalty by an environ-
mentally imposed factor has no theoretical barrier (Bressan
100
Fig. 2 Jatropha curcas planta-
tion. Plants are grown on waste
land under semi-arid conditions
(CSMCRI, Bhavnagar, India).
The insert shows a cluster of
fruits taken at the location
et al. 2001). Such approaches are currently being applied to
traditional crops (Bnziger et al. 2006) and to species that
are being promoted as bioenergy crops for biomass-derived
ethanol or biodiesel production, such as Miscanthus,
switchgrass or Jatropha, when grown under less than ideal
conditions (Heaton et al. 2008; Petyr et al. 2008).
There are two cardinal issues that demonstrate the
underlying necessity for an extremophile approach to
biomass and energy production. First, utilizing agricultural
land on which food can be grown productively interferes
with the imperative that we must produce enough food for a
growing human population. Second, land not covered by
forests and not useful for traditional crops without irrigation
amounts, astonishingly, to approximately one third of the
planets land mass. These savannahs, dry grasslands,
chaparrals and dry deserts, and likewise nutrient-poor
tropical forests, are increasingly under assault for agricul-
tural use. Rapidly increasing demands for food or industrial
materials from plants continues to increase pressure to
convert these marginally productive regions into areas for
crop production. Without political and technologically
sound alternatives this will happen because human pop-
ulations with no alternatives but to rely on subsistence
farming will be forced to take this route. The consequences
of intense individualized farming for food production with
crops that are not truly suitable in these areas are not only
discouraging but are potentially environmentally cata-
strophic, whether we focus on farming in the rainforests
of Indonesia, southeast Asia, the Amazon, or central Africa.
Clearly, a switch to bioenergy crops can free these human
populations from poverty and substantially help avoid the
danger of environmental collapse. Likewise, the arid zones
of sub-Saharan Africa, Australia, the southwestern United
States, western China, or the cold and dry vast regions of
central Asia and Siberia with their traditionally low
R.A. Bressan et al.
productivity become increasingly endangered by our
insistence on their use to alleviate inadequate agricultural
production in the present arable areas.
Conclusions
The worlds approximately 13 billion ha of land include
just over 10% that are highly suitable for agriculture.
However, the area suitable for marginal farming is maybe
three times that acreage (Food and Agriculture Organiza-
tion of the United Nations 2003). This less suitable farm
land, and to some variable degree all land, experiences
abiotic stresses that may become more pronounced, and
will likely become more erratic, in the wake of global
climate change. Deviation from the long-term mean of
temperature, wind and moisture for a particular area,
especially frequent and inconsistent variations, generates
most certainly the most serious challenge to sustaining crop
yield. As early as 1982, Boyer pointed to the high yield
reductions caused by abiotic stresses, and newer studies
have provided evidence that crop yields are still less than
one third of what could be achieved under ideal growth
conditions (Boyer 1982; Evans and Fischer 1999). A lucid
documentation of the constraints on productivity in African
and Asian farming systems documented the problems anew
(Waddington et al. 2010). These problems will persist at
least until extant crop species have been optimized to
present and new climates and only if we can add or uncover
genetic resources supporting higher and more stable
production in variable climates and tolerance of other
abiotic stresses within the genomes of our crops. There
is near universal consensus that drought is the most
damaging of the abiotic stresses that requires immediate
and determined attention.
Stress-adapted extremophiles provide energy without interference
Increasing irrigation, the remedy for combating drought
in past millennia, will not be practical or possible because
water is either not available or will become too energy
costly. Water will become less available for agriculture as
urbanization increases. Also, access to fresh water in desert
and semi-desert areas of the world is developing into a
political issue that could become explosive. Water scarcity
will also affect irrigation already in place and using less
water out of necessity will then accelerate soil salinization. As
the climate changes, additional challenges for agriculture will
focus on temperature extremes. Although some success in
protecting plants from drought and temperature damage have
recently been reported (Nelson et al. 2007; Rivero et al.
2007), overly optimistic predictions seem inappropriate
because the results so far indicate that fundamental solutions
to the drought problem are not yet at the implementation
stage.
No country is immune to climate change, but the
developing world will bear the brunt of the effects,
including some 7580% of the costs of the anticipated
damage (World Development Report 2010). As the
consequences of the likely changes come into clearer focus,
the results of inaction become equally clear, yet still the
most apparently efficient remedies are hotly debated. It
seems extremely important and prudent that food crop
productivity must be improved and that should not be
diminished by allocating arable land to biomass crops. In
light of this, crop improvement, whether by transgenic
means or based on molecular breeding advances, is finally
beginning to attach importance to crops that are better
protected against the influence of changing environmental
conditions (Nelson et al. 2007; Edmeades et al. 1999;
Bnziger et al. 2006; Moose and Mumm 2008; Messmer et
al. 2009; Tester and Langridge 2010).
The future we envision assumes that the immediate
use of wild, extremophile plants in small scale applica-
tions should go hand in hand with efforts to develop
agronomically a number of evolutionarily stress-adapted
species. These will strictly be reserved for energy
production on marginal soils, already degraded by
desperation agriculturists, with minimal impact on
important ecosystems. It will be essential that the ill-
adapted crop species presently used in such areas are
gradually replaced. Massive breeding efforts should be
devoted to develop EECs. Also, breakthroughs of the
recent past in genomics and improved understanding of
the regulation of gene and protein expressionincluding
epigenetic mechanisms that allow plants to respond in a
flexible way to environmental challenges (Chinnusamy
and Zhu 2009)must be employed. Genomics-based
selection approaches and transgenic modifications that
allow transfer of mechanisms from EECs into species with
high biomass productivity will be required.
101
Solar efficiencies will have the final word on the
problem. Solar energy irradiation of the earths surface
amounts to approximately 5.71024 J on an annual basis.
Photosynthetic organisms as a group utilize this energy in
fixing CO2 at ~21011 t/year (31021 J/year), whereas
amounts consumed by humans are ~31020 J/year (Hall
1979), representing only 10% of the energy converted
during photosynthesis. The typical photosynthesis efficien-
cy of plants is well below 1%. Compared to that value, the
efficiency of advance solar photovoltaic (PV) cells
approaches or exceeds 40%. Even the emerging PV cells
such as dye-sensitized or organic cells that could reason-
ably compete economically with current electricity produc-
tion show efficiencies higher than 6% at least for research
cells (Best Research-Cell Efficiencies, National Renewable
Energy Laboratory 2007). It seems clear that in the long
term utilization for primary energy production will move
away from bioconversions. We advocate that ethanol
production should be the first to find replacement, whereas
other bioenergies such as localized biogas generation can
be sustained. More efficient physical-chemical processes
and more direct solar energy-based sources will eventually
be required.
Biofuels have served as the primary energy source from
antiquity. All advances of human civilization have been
based on biofuels, as new sources were found and new
methods were developed for utilization. Non-biological
sources, such as photovoltaics, wind, wave, or geothermal
and other technological solutions are becoming competi-
tive. As a consequence, better engineered, efficient biofuel
resources will become integrated and adapted to particular
locations and conditions on a local scale. Bio-methane,
even bio-alcohols, will find their proper role in a future
economy designed to place prime importance on environ-
mental sustainability. In such a scenario the use of EECs
can provide significant input in a way that contributes to
food security and also protects important natural ecosys-
tems. However, implementation of these approaches should
be carefully restricted so as not to contribute or generate
additional population pressure, poverty or further income
imbalances, food shortage and excess, and land use and
abuse. This energy crop production should be viewed as
replacing poorly adapted cash crops or even subsistence
cropping where economically viable supply and marketing
infrastructures are available. Otherwise extremophiles that
provide specialty products such as high value oils, other
high energy products and localized bio-energy should be
grown. Increased water use and energy requirements also
require global economic and political adjustments; as
discussed by the tragedy of the commons parable,
solutions have no strictly scientific remedy (Hardin 1986).
Even for those who see the future with respect to energy
and food with less certainty, and perhaps with more
102
optimism, than our analysis suggests, a focus not just on the
immediate future but on the long run is of paramount
importance.
Epilogue
by a cherished friend and colleague, Lowell S. Hardin,
economist and architect in the debate leading to the
original green revolution (Meetings that changed the
worldBellagio 1969: The green revolution. Hardin
2008).
In the 1960s when the thrust to increase food crop
productivity in Asia was launched, serious scholars held
that famine was inevitable. They were wrong. The
production acceleration campaign succeeded. By 1975
India and Pakistan had increased food crop production so
greatly that they were self- sufficient. This unpredicted and
unprecedented phenomenon came to be known as the green
revolution. It was based on the application of strategies and
tactics that had earlier brought about the agricultural
revolution in the United States and Europe. While plant
breeding, and improved production technologies including
irrigation were involved, the green revolution had another
essential but little recognized component. That was incen-
tive policies formulated and administered by supportive
governments. These often included subsidies on inputs
(seed, fertilizer, irrigation water) and guaranteed prices for
the product (price supports, public acquisition programs).
Given todays even more complicated food and energy
security situation than existed during the food crisis of the
1960s, it is apparent that the strategies and tactics employed
to generate the green revolution are not in themselves equal
to the challenge. They need to be employed to their full
productive potential. But given the looming possibility (if
not probability) that the world may face a very tight supply
of crop land and water with an ever-tightening food and
energy situation, additional strategies such as the one set
forth here command accelerated investment in research and
testing. Determining the technological and economic
potential of generating energy from biomass produced with
stress-adapted extremophiles is an insurance strategy
helping make the future less insecure. For this to occur, as
with the green revolution, incentive policies (long-term
research support, public-private partnerships) must be
formulated, adopted and implemented.
The outline provided here points towards the use of
marginal lands and lands destroyed by irrigation over many
centuries for the cultivation of naturally stress-adapted
species, which could over time be converted into energy
crops. While the policies for such an undertaking are
missing, developing such policies, and their implementation
R.A. Bressan et al.
through long-term support by international organizations
and/or foundations can be imagined.
Acknowledgments Our work has been supported by funds from
King-Abdullah-University for Science and Technology of Saudi
Arabia, by the World Class University Program (Korea, R32
10148), by the Biogreen 21 Project of the Rural Development
Administration (Korea, 20070301034030), and by University of
Illinois and Purdue University institutional support.
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Spatari, S., Zhang, Y., & Maclean, H. L. (2005). Life cycle assessment Gujarat University, Ahmadabad,
of switchgrass- and corn stover-derived ethanol-fueled automo- India, in 1985. He worked as a
biles. Environmental Science & Technology, 39, 97509758. scientist at the Central Salt and
Tester, M., & Langridge, P. (2010). Breeding technologies to increase Marine Chemicals Research Insti-
crop production in a changing world. Science, 327, 818822. tute (CSIR) Bhavnagar, India,
USDA, NRCS. (2002). Plant Profile for Distichlis spicata (L.) Greene. 1987-2009, focusing on improv-
The PLANTS Database, Version 3.5 (http://plants.usda.gov/). ing stress tolerance and produc-
National Plant Data Center, Baton Rouge, LA 70874-4490 USA. tivity in crops and wild species,
Vincour, B., & Altman, A. (2005). Recent advances in engineering with emphasis on biofuel plants.
plant tolerance to abiotic stress: Achievements and limitations. He was a visiting scientist in the
Current Opinion in Biotechnology, 16, 123132. Department of Horticulture, Pur-
Waddington, S. R., Li, X., Dixon, J., Hyman, G., & de Vicente, M. C. (2010). due University, USA, 1996-97
Getting the focus right: production constraints for six major food crops and 1999-2000. He serves now
in Asian and African farming systems. Food Security, 2, 2748. as Director of Greenhouses, Plant Stress Genomics Research Center,
Westlake, D. F. (2008). Comparisons of plant productivity. Biological King Abdullah University of Science and Technology. His interest is crop
Reviews, 38, 385425. and plant improvement for abiotic stress tolerance and productivity.
Stress-adapted extremophiles provide energy without interference
Suk Ho Chung received his
B.S. degree in Mechanical
Engineering in 1976 from Seoul
National University and his
Ph.D. degree in Mechanical
Engineering in 1983 from
Northwestern University. He
worked as a professor in the
School of Mechanical and Aero-
space Engineering, Seoul Na-
tional University during 1984-
2009. He is now a professor in
Mechanical Engineering in King
Abdullah University of Science
and Technology, and Director of
the Clean Combustion Research Center. His research interests cover
combustion, pollutant formation, and laser diagnostics.
Dae-Jin Yun received his Ph.D.
degree in Agricultural Chemis-
try from Kyoto University,
Japan, in 1994. He worked as a
scientist at the Center for Plant
Environmental Stress Physiolo-
gy, Purdue University, USA,
1994-1998, focusing on stress
tolerance mechanisms in plant.
He has worked as a professor in
the Department of Biochemistry,
Gyeongsang National Universi-
ty since 1998. He serves now as
the Director of World Class
University program supported
by the Ministry of Education, Science and Technology, Korea. His
main research topic is to develop Plant Environmental Biotechnology
for Climate Change.
105
Lowell S. Hardin received his
Ph.D. from Cornell University,
USA, in 1943. During a long
career at Purdue, he has been a
teacher of farm management and
Head, Department of Agricultur-
al Economics professor, and is
now an emeritus in this depart-
ment. From 1965 to 1981 he
served as senior agriculturist
with the Ford Foundation. Dur-
ing his tenure there he became a
principal architect of the world-
wide system of thirteen interna-
tional agricultural research cen-
ters. His name is also associated with the discussions that initiated the
first green revolution in agriculture. He is a Fellow of the American
Agricultural Economics Association for which he has served as
secretary-treasurer, vice president and president.
Hans J. Bohnert received a
Ph.D. degree in General Biol-
ogy from the University of
Heidelberg, Germany, in 1972.
After working at the University of
Dsseldorf, Germany, CSIRO
Plant Industries, Australia, EMBL
Heidelberg and Max-Planck-
Institute for Plant Breeding, Ger-
many, he joined the Department of
Biochemistry, The University of
Arizona, USA, in 1983. In 2001
he moved to the University of
Illinois, Departments of Plant Bi-
ology and of Crop Sciences,
Urbana-Champaign, USA. He also serves as the director of the
Center for Comparative and Evolutionary Genomics. He is a
visiting professor at Gyeongsang National University, Jinju,
Korea. His interests center on understanding mechanisms confer-
ring abiotic stress tolerance in plants.