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 Chapter 12

Bioenergy

Daniel K. Y. Tan and Jeffrey S. Amthor

Additional information is available at the end of the chapter

[Link]

. Introduction

”ioenergy is the chemical energy contained in organic materials that can be converted into
direct, useful energy sources via biological including digestion of food , mechanical or
thermochemical processes [ ]. Over the past years, fossil fuel combustion provided the
energy for industrialisation and development of the modern economy. “round , total
energy consumption by humanity was about EJ yr [exajoules-
J see “ppendix for list
of abbreviations and units], mainly supplied by wood [ ]. ”y , the world total primary
energy supply was about EJ yr , or the equivalent of
-
million Mg of oil per year
Mtoe yr , which was almost double that of the
-
Mtoe supply in Figure [ ]. This
value is equivalent to global energy consumption by humans of TW × W , and
global energy demand is projected to increase to TW by [ ]. In , % of total energy
consumption was derived from fossil fuels, with only . % from renewable energy sources [ ].
Unfortunately, we are running out of fossil fuels, which originated from plant material
produced in ancient times, and combustion of these fossil fuels leads to emissions of CO and
the consequent global warming. Current proven reserves of oil would last only - years,
natural gas - years and coal - years at rates of consumption [ ]. “lthough some
experts claim that peak oil will occur in about years, others argue that the world is already
at peak oil production [ ]. In either case, fossil fuels are created at a slower rate than they are
now being consumed and cannot be considered as the world s main source of energy for more
than one or two more generations. This review gives an overview of the amount of energy that
can be harvested from the sun for contemporary biomass production, both for food and for
bioenergy.

© 2013 Tan and Amthor; licensee InTech. This is an open access article distributed under the terms of the
Creative Commons Attribution License ([Link] which permits
unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
300 Photosynthesis

Figure 1. World total primary energy supply from 1971 to 2009 by fuel (Mtoe) [3] *Other includes geothermal, solar,
wind, heat, etc.

. Photosynthesis as a source of food and bioenergy

Oxygenic photosynthesis occurs in cyanobacteria, algae and land plants and is summarised
by the equation

CO + H O + light energy ® éëCH O ùû + O

where [CH O] indicates a carbohydrate product of photosynthesis such as sucrose or starch.

Photosynthesis is the source of global food, feed, fibre and timber production as well as
biomass-based bioenergy. Each of these products of photosynthesis are renewable. Starch and
sucrose are the main products of photosynthesis and sucrose is the main form of carbon
translocated from leaves to other organs in plants.
The total energy from sunlight reaching the earth s surface is about TW[ ] nearly
EJ per year , about times the annual global human primary energy con‐
sumption of EJ [ ]. However, solar energy is geographically diffuse and this makes the
efficiency of conversion of sunlight important to capture this energy in useful forms. Due to
the Carnot limit, the maximum theoretically possible conversion efficiency for sunlight into
electricity is % [ ]. Photovoltaic cells have efficiencies around – % for converting
sunlight into electricity, but are limited to a maximum conversion efficiency of ~ % due to
the Shockley-Queisser limit [ ]. “ctual solar energy conversion by photosynthesis and
subsequent plant growth biomass production is much lower at around – % for productive
plant communities [ ].
 Bioenergy 301
[Link]

. Global primary production

Earth s land plants assimilate about Pmol of atmospheric CO each year, storing at least
temporarily nearly EJ yr in sucrose, starch and other carbohydrates. For marine
-

organisms the total may be - Pmol of CO yr- . Those are values of gross primary production
GPP or total photosynthesis. “nnual global net primary production NPP , defined as GPP
minus the respiration of the photosynthetic organisms, may be nearly Pmol C on land
equivalent to as much as EJ yr- and perhaps Pmol C in the ocean [ ]. NPP is critical
to present life on earth because it is the organic matter and associated energy potentially
available to all non-photosynthetic organisms for use in support of their growth, maintenance
and reproduction. NPP also contains the bioenergy potentially available to society.
Land plants may have an advantage over aquatic plants as they are able to photosynthesise
using leaves that can make use of the rapid diffusion of gases in air which is about times
faster than that in water [ - ]. Thus, cyanobacteria and algae in water may need to be well
stirred to support rapid photosynthesis and growth [ ]. Fortunately for oceanic photosyn‐
thesisers, surface waters are often vigorously mixed and where nutrients are available primary
production can proceed rapidly. Since oceanic NPP is dominated by phytoplankton, most of
the plant biomass there is photosynthetic as they do not have non-photosynthetic structures
such as roots and woody stems. Those oceanic primary producers represent only about . %
of global ocean + land primary producer biomass due to rapid turnover time in the oceans
average to days compared to much slower turnover on land average of about years
[ ]. Large ocean area provides a significant potential for biomass production, though nutrients
are often limiting and harvesting oceanic biomass is difficult and challenging. ”ecause of
relative ease of harvesting, and the longer life time of land plants, nearly all current bioenergy
harvesting is from terrestrial plants.

. Converting solar energy to biomass

To describe the component processes associated with the use of solar energy to produce
biomass, Monteith s equation [ ] can be used

Y = . ´ St ´ e i ´ e c ´ e p

Where Y is biomass energy yield J m- of ground St J m- is the total incident solar radiation
during the growing season ฀i is light interception efficiency fraction of incident radiation
absorbed by a plant s photosynthetic apparatus, J J- ฀c is photosynthetic conversion efficien‐
cy, including metabolic costs of growing new biomass from products of photosynthesis J J-
in resulting biomass and ฀p is partitioning efficiency or harvest index J J- .
Photosynthetically active radiation P“R is approximately confined to the - nm
waveband [ ] which contains about % of total solar energy reaching Earth s surface St [ ].
302 Photosynthesis

Thus, about half of the incident solar energy is unavailable to higher-plant photosynthesis,
which is accounted for in the coefficient . in the equation above see Figure . In addition,
the fraction of solar radiation absorbed by plants or ฀i, depends on leaf area and orientation.
“ full canopy can potentially absorb about % of incident P“R with perhaps % of that
absorption associated with chloroplasts [ ]. Partitioning efficiency ฀p or harvest index is the
amount of total biomass energy partitioned into the harvested portion of the crop for a biomass
crop that may approach %, but for a seed crop can be as low as %. The amount of energy
in a unit mass of plant material also varies, being about - MJ kg- for typical biomass, but
as much as - MJ kg- for oilseeds [ , ]. During the Green Revolution, the dwarfing of
the crop-plant stem improved partitioning efficiency ฀p [ ] and selection of larger-leaved
cultivars improved light interception efficiency ฀i , but there has been little apparent im‐
provement in photosynthetic conversion efficiency ฀c .

. Potential and actual photosynthetic conversion efficiency

The observed minimum quantum requirement of - mol photons per mol CO assimliated
in C photosynthesis represents an absolute limit on biofuel production from sunlight, in spite
of claims for biomass production usually by algal systems that would correspond to signif‐
icantly smaller quantum requirements [ ]. That range corresponds to C photosynthesis in
the absence of photorespiration, which in the current atmosphere increases minimum quan‐
tum requirement to about mol mol- . ”ut due to light saturation, and other factors below ,
biomass production, especially over an annual cycle, cannot approach limits set solely by
minimum quantum requirements [ , ].
The potential maximum efficiency of converting solar energy to biomass energy is estimated
at about . % for algae [ , ], . - . % for C land plants and . - . % for C land plants at
- ˚C and present atmospheric [CO ] see Figure [ , ]. C plants can be more efficient
than C plants because they are able to suppress photorespiration through a combination of
biochemical and anatomical innovations that arose relatively recently in plant evolution. These
innovations presumably were a response to declining global atmospheric [CO ] during the
past million years.
“ctual maximum conversion efficiency is generally lower than the calculated potential
efficiency at around . % for algae [ ], . % and . % for C and C crops [ ], respectively,
across a full growing season see Figure due to insufficient capacity to utilise all radiation
incident on a leaf, and photoprotective mechanisms that impair efficiency. The actual photo‐
synthetic efficiency of mature C forest stands was also calculated to be between . to . %
[ ]. Of course, plants are self-regenerating and self-maintaining whereas photovoltaic cells
are not.
The low yields from biomass energy production are frustrating compared with photovoltaic
cells that have efficiencies of up to %, and this is due to the following limitations in plants
[ , , , , ]

. Two photosystems Photosystems I and II in series

 Bioenergy 303
[Link]

. Dependence on photons limited to the approximate waveband – nm

. Inherent inefficiences of enzymes and biochemical processes

. Light saturation under bright conditions and associated photoinhibition in Photosystem

II

. Respiration, an absolutely essential process for life and growth [ ], which consumes
- % of the energy contained in the products of photosynthesis and

. Plants are living organisms that spend about half of each day in the dark, when they need
to use previously generated carbohydrate stores to keep themselves metabolically active
and growing.

Figure 2. Minimum energy losses associated with biomass production. Wedges show the percentage of energy from
solar radiation remaining (inside arrows) and percentage losses (at right) from an original 100% calculated for several
stages of photosynthetic and biosynthetic energy transduction from sunlight incident on a plant community to new
plant biomass [18]. This analysis indicates that a theoretical maximal photosynthetic energy conversion efficiency is
4.6% for C3 and 6.0% for C4 plants at 25-30 oC.
304 Photosynthesis

(a) Potential (maximal) efficiency

100solar energy input (incident)

55.6not photosynthetically active (incident)
44.4photosynthetically active (incident)
3.7canopy albedo
40.7absorbed by leaves
4.44inactive absorption
36.26absorbed by photosynthetic pigments
28.34lost as heat
7.92assimilated in photosynthesis
1.98respiration
5.94new phytomass

(b) Actual (observed) efficiency

100solar energy input (incident)

55not photosynthetically active (incident)
45photosynthetically active (incident)
7canopy albedo
38absorbed by leaves
3inactive absorption
35absorbed by photosynthetic pigments
31lost as heat
4.0assimilated in photosynthesis
1.0maintenance respiration
3.0available for growth
0.8growth respiration
2.2new phytomass

Figure 3. estimates of (a) potential efficiency (theoretically maximal) and (b) actually efficiency for biomass production
of a healthy crop [17]. These results are still generally applicable.

The current bioenergy enterprises are focussing on C crops such as sugarcane, maize, sweet
sorghum, switchgrass and miscanthus presumably due to the higher energy conversion
efficiency. However, this advantage of C over C will disappear as atmospheric [CO ]
approaches µL L- .
 Bioenergy 305
[Link]

. Improving energy conversion efficiency

Due to the apparently low actual energy conversion efficiency in whole-plant photosynthesis
i.e., - % , much discussion has focused on improving photosynthesis to improve crop yield
potential including [ , , , ]
. Engineering C crops to use C photosynthesis. This would potentially suppress photo‐
respiration and increase net photosynthetic efficiency by as much as % and increase
both water and nitrogen use efficiencies [ , ]. There is an ongoing ambitious research
program, led by the International Rice Research Institute IRRI , to convert the normally
C rice to a C system by transforming rice to express Kranz anatomy and the C metabolic
enzymes [ ].
. Improving both rubisco s catalytic rate of carboxylation kcat and specificity for CO
relative to O τ . This would improve the efficiency of rubisco as a catalyst of CO
assimilation [ , ]. Unfortunately, τ and kcat are inversely related across many rubiscos
found in nature [ ]. “nother complication for engineering an improved rubisco is that it
is composed of eight large chloroplast-encoded subunits and eight small nuclear-encoded
subunits, and assembling modified subunits in chloroplasts remains a challenge [ ].
. Minimising, or truncating, the chlorophyll antenna size of chloroplast photosystems. This
would potentially improve solar conversion efficiency by up to -fold in high light, which
normally saturates photosynthesis [ ]. Individual cells or chloroplasts would have a
reduced probability of absorbing sunlight, allowing greater transmission to leaves lower
in a canopy and a more uniform distribution of light across leaves within a canopy, hence
reducing dissipation and loss of excess photons in non-photochemical quenching
NPQ .
. Improving the recovery rate from the photoprotected state. This would potentially
increase carbon uptake by crop canopies in the field [ ]. The xanthophyll photoprotection
system protects plants from damage from absorption of excess light the reduction of
photosynthesis by dissipation of photons by NPQ . High-yielding rice are reported to
recover more quickly from photoinhibition than traditional varieties [ ].

. C photosynthesis

C crops include wheat, rice, cotton, barley, soybean, bean, chickpea, algae, palm and peanut.
C photosynthesis of CO forming fructose -P can be summarised by

CO + “TP + N“DPH ® fructose -P + “DP + Pi + N“DP

In principle, the “TP and N“DPH required to assimilate one CO molecule can be produced
during absorption of eight photons of P“R. ”ecause mol photons P“R contain on average
306 Photosynthesis

. MJ, and because about . MJ of energy per mol C is stored in carbohydrates, the potential
efficiency of converting absorbed P“R to biomass approaches % for C photosynthesis or
about % for total solar radiation . That efficiency occurs only in low light, however under
a bright sun, C photosynthesis becomes light-saturated. In addition, the process of photores‐
piration, which is relatively rapid in C plants, especially at higher temperature, is a significant
constraint on CO assimilation in C plants. “s much as one third of the C assimilated in C
photosynthesis can be almost immediately lost to photorespiration with present atmospheric
CO concentration higher CO concentration not only stimulates photosynthesis, but inhibits
photorespiration . In sum, at about °C, the efficiency of converting absorbed P“R into
carbohydrate may be about % in C plants, accounting for photorespiration, but ignoring
light saturation. Moreover, that efficiency does not account for plant respiration, and some
respiration is essential for growth and maintenance processes.

. C photosynthesis

C plants include maize, sugarcane, sorghum, millet, miscanthus and switchgrass. The C
system involves the specialised metabolism and Kranz leaf anatomy to concentrate CO in the
bundle sheath cells. Normal C photosynthesis takes place in the bundle sheath cells in C
plants, but because the CO concentration there is quite high, photorespiration is greatly
suppressed. The C cycle, which concentrates the CO in bundle sheath cells, requires two “TP
to assimilate a CO in the mesophyll, release it in the bundle sheath and regenerate the CO
acceptor in the mesophyll. Some CO leakage from the bundle sheath is inevitable, and this
requires that the C cycle operates more quickly than the C cycle in C plants. Hence, C
photosynthesis may require at least . “TP CO - more than C photosynthesis, based on a
modest CO leak rate of % from bundle sheath cells [ ]. In spite of the extra energy cost of
the C cycle, C photosynthesis responds better to bright sunlight and to higher temperatures
than C photosynthesis because of suppressed photorespiration. “t cooler temperatures e.g.,
- oC , however, C photosynthesis is superior because photorespiration operates slowly at
low temperature. In addition, many C plants are sensitive to low temperature. The C plant
miscanthus Miscanthus × giganteus is relatively tolerant of low temperature, and it may be a
good source of germplasm for improving the low temperature tolerance of other C plants [ ].
In terms of efficiency, C photosynthesis might retain as much as - % of the energy in
absorbed P“R in carbohydrate products, again before any required respiration and ignoring
light-saturation. That efficiency is relatively insensitive to temperature, at least over the normal
range experienced by typical C crops during daylight hours.

. CAM photosynthesis

Commonly cultivated C“M crassulacean acid metabolism plants include agave Agave spp. ,
Opuntia Opuntia spp. , pineapple Ananas comosus , Aloe vera, and vanilla Vanilla planifolia .
C“M plants are well adapted to arid and semi-arid habitats. They open their stomata at night
 Bioenergy 307
[Link]

and take up CO in the dark to form malic acid, which is then metabolised to release CO for
photosynthesis during the following day, but with their stomata closed [ , ]. ”y closing the
stomata during the day, less water is lost, resulting in high water use efficiencies with a trade-
off of lower growth rates. C“M imposes an additional metabolic cost of ~ % compared with
the standard C pathway due to the transport of malic acid into the vacuole at night and
conversion of C residue back to the level of storage carbohydrate during the daytime [ ].
C“M plants have been suggested to have potential for food, fibre and biofuel production in
dry marginal lands [ , ].

. World food energy demand

The energy contained in food consumed per person is only about MJ day- equivalent to
kcal per day, ”tu or W [ ]. Hence, the food energy needed to feed the world s
current seven billion persons is ~ EJ yr- , which is only about % of the world s ~ EJ of
annual energy consumption, but more than % of global land NPP. The world s food
production system consumes about EJ yr- and hence, it takes about units of fossil energy
to produce unit of food energy [ ]. In the United States, the overall energy input/food output
ratio is even larger, around to [ ]. Most of the energy consumption ~ % occurs after the
farm gate, during transportation, processing and retail. Globally, one third of food, around .
billion Mg, is discarded including spoilage each year, and a similar share of the total energy
inputs are embedded in these losses [ ].

Global population is projected to increase to - billion within - years [ ]. In developing

countries, food consumption per person is rising with increased consumption of animal
protein with the livestock revolution [ ]. “verage annual meat consumption is projected to
rise from kg person- in to kg person- by [ ]. Grazing livestock already occupy
a quarter of the world s land surface, and the production of livestock feed uses a third of arable
cropland [ ]. With future increases in global population and per capita food consumption,
global food production will have to increase by as much as % to meet the increased demand
in , an annual growth rate in food supply of . % yr- [ ]. In principle, this means that by
the energy consumption for global food production may increase by EJ yr- from
today s EJ yr assuming the energy conversion efficiency remains constant.
-

In ancient civilisations, most of the energy used for farming was provided by animals and the
nutrients were derived from animal manure. During and after the Green Revolution, depend‐
ence on non-renewable fossil fuels resulted in a conversion of fossil energy into food energy,
but in an inefficient way. “griculture uses about % of the global fossil fuel energy of which
% is used for the production of nitrogen fertiliser from natural gas and atmospheric N using
the Haber-”osch process [ ] with a stoichiometry of about MJ kg- N [ ]. The dependency
of agriculture on fossil fuels has resulted in commodity food prices being closely linked with
global energy prices [ ]. Hence, food prices tend to fluctuate and trend upwards in parallel
with energy prices. It is instructive to compare maize production in Mexico using human
labour with a hoe and sickle returning . times as much energy in the harvested crop as
308 Photosynthesis

used in production of that crop with a return of less than times for mechanised maize
production in the United States Table . The U.S. crop was, however, more than nine times
as productive.

Crop Country Tillage Yield Inputs Output Energy ratio

(Mg ha )-1
(MJ ha ) -1
(MJ ha ) -1

Groundnut Thailand Buffalo 1.28 8 048 20 892 2.60

Groundnut USA Mechanised 3.72 45 817 64 051 1.40

Maize Mexico Human 1.94 2 687 28 881 10.70

Maize Mexico Oxen 0.94 3 222 13 982 4.34

Maize USA Mechanised 8.66 33 961 130 396 3.84

Rice Borneo Human 2.02 4 327 30 626 7.08

Rice Philippiines Carabou 1.65 7 638 25 126 3.29

Rice Japan Mechanised 6.33 34 405 96 163 2.80

Rice USA Mechanised 7.37 49 542 110 995 2.24

Soybean USA Mechanised 2.67 12 609 40 197 3.19

Wheat USA Mechanised 2.67 17 740 35 354 2.13

Table 1. Energy use in grain and legume production [49].

In developing countries, populations tend to have a cereal-based diet and are effectively at a
lower trophic level in the food chain, while populations in developed countries tend to
consume more meat and operate at a higher trophic level. Production of livestock, on average,
may require kg of wheat for the production of kg of meat [ , ]. Therefore, in developed
countries where kg of cereal and kg of meat are consumed per year, the total need for
food and feed is kg of cereal per person per year [ ]. Overfishing of the ocean predators
e.g., killer whales, tuna, salmon at high trophic levels has also led to the decline in ocean
fisheries yield [ ]. It is important that cereal crops supply % of the calories consumed by
humans on the global scale with the remainder supplied by potatoes, beans and other crops,
with marine animals now contributing only % of the human food supply [ ]. To increase the
energy efficiency of our primary food production system, we should focus on primary
production in agriculture e.g., cereals and aquaculture e.g., algae, phytoplankton rather
than secondary production e.g., livestock, fish .

. Biofuels

In addition to providing food and feed, plants are an important source of fuel. Indeed, biofuels
are not a new concept. In ”.C., the Syrian city of “ntioch had public street lighting fuelled
by olive oil. More recently, the German inventor Rudolph Diesel demonstrated his engine that
ran on peanut oil at the World Fair in Paris. In simple terms, the nearly EJ contained
 Bioenergy 309
[Link]

in annual global NPP is about times current global energy demand ~ EJ [ ]. That NPP,
however, includes vast amounts of biomass that cannot be physically or economically
harvested including national parks . In , biomass, including agricultural and forest
products and organic wastes and residues, accounted for nearly % of the world s total
primary energy supply [ ], with fraction less than % in developed countries, but as high as
- % in developing countries [ ]. Replacing fossil fuels with renewable energy sources
derived from sunlight, such as solar, hydro or biomass is very challenging as these energy
sources have a lower energy density than fossil fuels and are generally more expensive [ ].
In some developing countries, as much as % of total energy consumption is supplied by
biomass [ ]. Solid biomass such as firewood, charcoal and animal dung represent up to %
of all biofuels [ ].
Since the beginning of civilisation, humans have depended on biomass for cooking and
heating, and many developing countries in “sia and “frica are still dependent on traditional
sources of biomass. Liquid biofuels account for only % of total bioenergy, and they are mainly
significant in the transportation sector. Transportation accounts for % of global energy
consumption and % of global oil production, and liquid biofuels supplied only % of total
transport fuel consumption in [ ]. The automative industry currently uses relatively
energy inefficient internal combustion engines to burn liquid fuels e.g., gasoline and diesel .
Electric car motors have a . times higher energy efficiency than internal combustion engines,
but the lightness and compactness of liquid fuels still have a fifty-fold higher energy storage
than the best available batteries [ ]. Hydrogen fuel cells may replace electric motors in the
future but this is still in the developmental phase. In the meantime, liquid biofuels are the
transition renewable alternative to fossil fuels. Globally, liquid biofuels can generally be
classified into three production sources maize ethanol from the United States, sugarcane
ethanol from ”razil and rapeseed biodiesel from the European Union. In , ”razil and the
United States produced % of the billion L of global bioethanol and the European Union
produced % of the billion L of global biodiesel [ ]. For the rest of this chapter, we use
the term biofuels to refer to liquid biofuels. First generation biofuels refer to the traditional or
conventional supply chains based on food crops, whereas second generation biofuels require
more complex and expensive processes and are generally operating in pilot plants and not yet
widely available on the market.

. First generation biofuels

The first generation of biofuels is produced from starches, sugars and oils of agricultural food
crops, including maize, sugarcane, rapeseed and soybean. Carbohydrates are fermented to
bioethanol, which is mixed with gasoline as a transporation fuel. ”ioethanol, produced mainly
from sugarcane, replaced % of gasoline used in ”razil in , with the introduction of flex-
fuel vehicles allowing high-blending of bioethanol with petrol all petrol blends in ”razil
contain % bioethanol [ ]. In the United States, up to % of the maize crop was used for
bioethanol production in . If all the main cereal and sugar crops wheat, rice, maize,
sorghum, sugar cane, cassava and sugar beet representing % of global cropland were to be
hypothetically converted to ethanol, this would correspond to only % of total petrol use in
[ ], and would leave no cereals or sugar for human consumption, although the reduced
sugar in the human diet would have health benefits. Oils/fats i.e., a mixture of triglycerides,
310 Photosynthesis

free fatty acids, and/or phospholipids are converted to biodiesels, potentially competing with
food and feed production from oilseed crops such as rapeseed including canola and soybean.
”iodiesel, a supplement or replacement to traditional diesel, is also produced from animal fats
tallow .

. Second generation and advanced biofuels

Due to food and energy security concerns, many countries are promoting bioenergy crops that
can be grown on land not suited for food production, so that the two systems are complemen‐
tary rather than competitive [ , ]. Second generation biofuels refer to the range of feedstocks
e.g., dedicated energy crops such as miscanthus, switchgrass, jatropha, pongamia, agave,
Indian mustard, sweet sorghum, algae, carbon waste , conversion technologies e.g., fast
pyrolysis and supercritical water , and refining technologies e.g., thermo-chemical Fischer-
Tropsch methods used to convert biomass into useful fuels Figure [ ]. There is a fine line
between a first and second generation biofuel. For example, sugarcane is a first generation
biofuel feedstock sucrose but co-generation for electricity using sugarcane residue bagasse
as a fuel is also possible, and sugarcane residues may serve as future feedstocks in second
generation ligno-cellulosic bioethanol production [ ]. Ligno-cellulosic bioethanol is based on
the conversion of lignocellulosic compounds, made up of chains of about glucose and
other small organic molecules, into sugars with sophisticated methods of acid or enzymatic
hydrolysis. Those sugars can then be converted to fuel using tradiational methods. This means
that non-food products such as cereal and wood residues can be converted to ethanol instead
of remaining as a waste by-product. These lignocellulosic residues are mainly cell walls that
make up - % and - % of the stems of woody and herbaceous plants, respectively, and
about - % in their leaves, and consists of around - % cellulose, - % hemicelluloses
and - % lignins [ ]. There are a few examples of commercial ligno-cellulosic plants. For
example, Swiss company Clariant opened a ligno-cellulosic plant in Germany in that can
produce up to Mg of cellulosic ethanol from Mg of wheat straw [ ]. Where lignin
cannot be converted to small sugars easily through biochemical processes, it can be burnt for
co-generation of bio-electricity.

“nother potential bioethanol feedstock is agave Agave spp. which is adapted to semi-arid
land unsuitable for food production [ , ]. “gaves are well-suited for biofuel production as
they can be grown in sandy soil with little or no irrigation and are less likely to be weedy.
“gave have above-ground productivities similar to that of the most efficient C and C crops
- Mg ha- yr- dry biomass , but with only % of the water required for cultivation [ ].

Sisal Agave sisalana is mainly produced in east “frican countries of Kenya, and Tanzania, as
well as in ”razil, China and Madagascar. The sisal leaf contains about % by weight of
extractable hard fibre vascular tissue , the remaining % being water and soluble sugars
which is disposed of during the decortication process into rivers and the sea, causing pollution,
eutrophication and water contamination [ ]. Production of ethaonol and bioenergy from sisal
juice from the sisal leaves and stems is under pilot testing at the Institute for Production
Innovation at the Uninversity of Dar es Salaam and “alborg University [ ]. The first field
experiment of blue agave Agave tequilana as a biofuel crop was planted in in the ”urdekin
 Bioenergy 311
[Link]

River Irrigation “rea of Queensland, “ustralia [ ]. ”lue agave can acheive strong growth
rates by potentially switching from C“M to C photosynthesis if there is sufficient water
supply [ ]. “pproximately . Mha of arid land was used to grow sisal for coarse fibres sisal
but this has fallen out of production or abandoned due to competition with synthetic fibre [ ].
In theory, this crop area . Mha alone could provide . billion L of ethanol if agave were
re-established as a biofuel feedstock without causing indirect land use change [ ].

In the meantime, new and novel feedstock conversion technologies are being developed such
as fast pyrolysis and supercritical water treatment that can now convert nearly any biomass
feedstock, such as wood residues, agricultural residues e.g., wheat and maize stalks , woody
plants, and C grasses [e.g., switchgrass Panicum virgatum , miscanthus and sweet sorghum]
into a green biocrude that can be processed into jet fuel, biodiesel, or bioethanol [ ]. Hydrogen
H is designated as a third generation biofuel, when it is produced from biomass by algae or
enzymes [ ]. H is a fuel whose combustion produces only water, although future technological
breakthroughs are needed before H can be produced economically.

Figure 4. The advanced biofuels value chain [60]. *Conversion technologies include fast pyrolysis and supercritical wa‐
ter treatment that transform feedstock into “green crude“ which is similar to crude oil. ** Macroalgae are multicellular
organisms (seaweeds) with low lipid content but are high in carbohydrates. *** Supercritical water treatment is a ther‐
mochemical process which involves subjecting the biomass to controlled temperature and pressure conditions in the
presence of appropriate catalysts to produce a “green crude“.
312 Photosynthesis

There is also a move to source oilseed from non-food dedicated energy crops grown on
marginal land. These crops might include jatropha Jatropha curcas , pongamia Millettia
pinnata , Indian mustard Brassica juncea , and microalgae. The recent failure of jatropha as an
energy crop in India and other developing countries due to a lack of bioenergy policy high‐
lights the need for investment in research and policy development before starting on large-
scale investments [ ]. Pongamia is a tropical tree legume Fabaceae family and is a native of
India and northern “ustralia. It has been used as a biofuel crop in India for some time, and is
well-suited to marginal land as it is regarded as both a saline- and drought-tolerant species.
The seeds contain about % extractable oil, predominantly in the form of triglycerides is rich
in C fatty acid oleic acid and has relatively low amounts of palmitic and stearic acid,
making it useful for the manufacture of biodiesel [ ]. In India, the de-oiled cake of pongamia
i.e., the leftover component of seeds following solvent extraction, and containing up to %
protein is used as a feed supplement for cattle, sheep, and poultry [ ]. Opportunities exist
for a sustainable pongamia agroforestry program to supply biodiesel in northern “ustralia,
although substantial infrastructure investment in processing plants would be needed [ ].
Indian mustard is another potential annual oilseed crop being developed in India and
“ustralia. It is drought-tolerant annual rainfall - mm and many varieties can express
greater osmotic adjustment than canola [ ]. Indian mustard was up to % more productive
than canola under dry conditions, but not under normal rainfall conditions in northwest New
South Wales, “ustralia [ ]. “n Indian mustard breeding program for biodiesel production
was commenced in at the University of Sydney s I.“. Watson Grains Research Centre at
Narrabri, “ustralia. Indian mustard is now part of a four year rotation at the Watson Centre.

Microalgae can be cultivated in open raceway ponds or closed photobioreactors, harvested,

extracted and then converted into a suitable biofuel such as biodiesel. Raceway ponds are
shallow no more than cm deep raceways and contents are cycled continuously around the
pond circuit using a paddlewheel. Most commercial algal producers are currently using open
raceway ponds as these require lower capital costs to set up but may result in increased
evaporation and risks of contamination [ ]. Photobioreactors are closed systems which offer
better control over contamination and evaporation but have higher capital and operating costs
than open raceway ponds [ ]. The surface area/volume ratios of photobioreactors are also
almost double that of the open pond, hence doubling the energy recovered as biomass and
potential productivity [ ]. Surface fouling due to competitors e.g., other algae , grazers and
pathogens e.g., bacteria are a major problem with photobioreactors and cleaning can be a
major design and operational problem [ ].

Despite the development of microalgae as a feedstock for biodiesel production, there are
problems scaling up from laboratories to commercial production [ , ]. Key limitations to
algal production in raceways or photobioreactors are the need for stirring, provision of
nutrients for optimal growth and very large surface areas required to capture significant
amounts of sunlight [ ]. Other problems are pathogen attack, ageing of algal cultures, and
lack of system optimisation [ ]. The need to de-water and dry the algal biomass can consume
up to % of the energy input of the process [ ]. Despite their potential productivity per unit
surface area, and containing up to % lipids as storage products, algal biodiesel is not yet
economically competitive with petroleum diesel algal diesel was recently priced at USD .
kg- compared with petroleum-based diesel at USD . kg- [ ].
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. Lifecycle analysis and energy balance

Life cycle analysis LC“ is a tool to take into account the inputs and outputs of a food or
biofuel crop production system, including the growing of the crop and its subsequent proc‐
essing the technique is also used to assess the energy efficiency and impact of food and biofuel
crops on greenhouse gases [ ]. Ecologists can relate an LC“ to a foodweb or ecosystem model
that traces the fluxes of energy through the system. Net energy value NEV is an efficiency
term calculated as the difference between the usable energy produced from a crop and the
amount of energy required for the production of that crop [ ].

Three annual crop management systems, conventional several tillage operations for weed
control, seedbed preparation, seeding , conservation reduced, minimum and no-till systems ,
and organic intensive tillage for seeding, weed control were compared in Canada and Spain
[ - ]. Generally, energy inputs for the conservation system were % lower than for the
conventional system due to lower fuel and machinery use from reduced tillage [ ]. How‐
ever, fertiliser and pesticide rates were often increased in response to increased soil water,
resulting in a similar total energy use by conventional compared with conservation systems
[ - ]. In contrast, there was a reduction in energy input in organic systems due to the use of
organic fertilisers instead of synthetic pesticides and fertilisers [ ]. In terms of energy
output input ratio, organic farming in Spain was . times more energetically efficient .
than either the conventional or conservation systems . and . , respectively [ ].
Inclusion of a leguminous forage crop e.g., vetch, chickpea into canola and cereal e.g., wheat,
barley rotations increased the energy efficiency and output under all management systems
[ , ]. Legume-rhizobial associations are effective solar-energy-driven systems fixing
atmospheric N into ammonia with minimal CO emissions compared to industrial nitrogen-
fertiliser production. Legumes fix nitrogen and thus reduce synthetic N fertiliser use in farming
systems they also enhance the productivity of subsequent crops through breaks in the disease
cycle [ ]. Pulses contribute about million Mg of fixed-N per year globally, accounting for
one third of the toal biological N fixation in agroecosystems [ ].

The energy efficiency of biofuels can also be termed the fossil energy ratio FER expressed as
the ratio of the amount of fuel energy produced to the amount of fossil fuel energy required
for that production [ ]. “n FER < indicates a net energy loss, whereas an FER > represents
a net energy gain. Life cycle assessments for biofuels have also shown that ”razilian sugarcane,
agave, and switchgrass ethanol could achieve positive energy balances and substantial
greenhouse gas offsets, while maize in the United States and China offers modest or no offsets
[ ]. The bioenergy created in sugarcane and agave ethanol, and in palm oil, is at least four
times the amount required to produce it, while maize in the United States and China release
almost as much energy when they are burnt as the energy that is consumed in growing and
processing them Figures and [ , ]. Sugar crops usually produce more ethanol per ha
with a better energy balance than starch crops because sugar crops produce higher sugar
amounts per ha than starch crops and sugar sucrose can be directly fermented, whereas
starch polymers have to be hydrolysed before being fermented by yeast [ ]. In general the
energy gain and conversion of solar energy into biomass in the sub-tropics is substantially
greater than any achievable in temperate zones [ ], possibly due to the longer growing season
and higher levels of solar energy over an annual cycle. For example, the FER of sugarcane in
”razil was . - in , compared with . for maize-ethanol in the United States and . for
314 Photosynthesis

sugarbeet in Europe [ ]. There is already evidence of a land-grab with countries e.g., China
and the Middle East securing their own energy and food security by acquiring large areas of
subtropical land in “frica and “sia [ ]. Many countries may never be able to establish a
position of energy or food independence or anywhere near approaching it [ ]. For example,
Sweden is importing ”razilian bioethanol as its main source of renewable transportation
energy, due to the climatic constraints of growing biomass for liquid fuels within Sweden [ ].
FER of microalgal-based Chlorella vulgaris biodiesel produced in raceways is . , which is
. times as energy intensive as conventional diesel FER of . ] in the United States [ ]. This
current negative energy balance is unacceptable unless the production chain can be fully
optimised with heating and electricity inputs decarbonised [ ].

Figure 5. Estimated ranges of fossil energy ratio (FER) of selected fuel types [54, 86]. Note: The ratios for cellulosic
biofuels are theoretical.
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Figure 6. Energy input and output per MJ ethanol produced for various feedstocks including Mexican agave, Brazilian
sugarcane, and United States maize (corn) and switchgrass [64]. Abbreviation: DDGS, Dried Distillers Grains with Solu‐
bles.

. Carbon footprint of food and biofuels

The world s food production system is inefficient, globally taking four units of energy to
produce one unit of food energy. Globally, agriculture also accounts for ~ % of nitrous oxide
N O and % of methane CH emissions [ ]. On-farm, N O emissions are mainly associated
with the use of nitrogenous fertiliser, while CH emissions are mainly from digestion in
ruminant livestock e.g., cattle and sheep . The carbon footprint is the total amount of green‐
house gases GHG associated with the production, processing and distribution of food and
biofuel crops expressed in carbon dioxide equivalents CO e . This can range from . to .
kg CO e per kg of wheat in New South Wales, “ustralia [ ], to . kg CO e per kg of wheat
in Canada [ ]. In Canada, legumes like chickpea, dry pea, and lentil may emit only . to
. kg CO e per kg of grain, about half that of wheat, due to biological N fixation by the
legumes. “s with the energy balance, durum wheat emitted % lower CO e when preceded
by an N-fixing crop, compared to when the crop was preceded by a cereal, highlighting the
benefits of including a legume in a cereal rotation on GHG balance [ ].
Since nearly % of the energy used in the food supply chain is in the postharvest phase [ ],
there is a fashionable trend for an eat-local movement to reduce `food miles`. Food miles refers
to the distance a food commodity travels from the point of production to the point of con‐
sumption and the related energy and CO emitted during transportation [ ]. However,
shipping food for long distances may sometimes require less energy and emit less CO . For
example, even when shipping was accounted for, New Zealand dairy products imported into
the UK used half the energy of their UK counterparts, and in the case of lamb, a quarter of the
energy due to grass-fed conditions in New Zealand compared with the energy-intensive
316 Photosynthesis

system used in the UK [ ]. New Zealand was % more energy efficient for apples, and the
energy costs of shipping of onions was less than the cost of storage in the UK, making New
Zealand onions more energy efficient overall [ ].
Over % of the food energy is lost through wastage in both developed and developing
countries [ ]. In developing countries, food is mainly lost due to pests, and spoilage due to
the lack of cold storage and food-chain infrastructure. In developed countries, food safety
issues have resulted in the over-reliance of `use by` dates resulting in good food being
discarded in landfills instead of being used, e.g., for animal feed or compost [ ]. This wastage
can be reduced through improved education, better legislation and research in postharvest
technology to reduce food wastage. We can invest in better diagnostics that monitor food
spoilage such as temperature- and time-sensitive inks on food package that cause labels to
change colour if the food has been exposed to the wrong temperature for too long [ ].
Restaurants can stop serving super-sized portions.
GHG emissions during agricultural production of biofuel crops contribute - % to the GHG
balance of maize ethanol in the United States [ ] and more than % in pure vegetable oils [ ].
In theory, biofuel feedstocks remove CO from the air and can potentially reduce greenhouse
gas emissions. However, clearing of undisturbed native ecosystems such as rainforest and
savanna or grassland for biofuel production also increases net GHG production due to change
in land use [ ]. For example, a hectare of maize grown for bioethanol can sequester . Mg
ha- yr- of CO e, but each hectare of forest converted to maize field has up-front emissions of
- Mg CO e and each hectare of grassland converted to crop releases - Mg CO e
equivalents. Hence, maize-based bioethanol production might double GHG emissions over
years and increase GHGs for years [ ]. Converting lowland tropical rainforest in Indo‐
nesia and Malaysia to oil palm biodiesel crops would result in a carbon debt of Mg CO
ha . This might take - years to repay, while sugarcane bioethanol produced on ”razilian
-

Cerrado woodland-savanna might take years to repay [ ].

. Water footprint of food and biofuel

“griculture accounts for about % of global fresh water consumption [ ]. Energy is also
needed to pump water for irrigation. The water footprint WF of a product, such as food or
biofuel, is the total volume of fresh water used for production and processing, through to
eventual use of the the product [ ]. In general, the WF of biofuels is up to - times larger
than the WF of fossil fuels [ ]. For example, water consumption of bioethanol processed
from rainfed maize grain is . L km- travelled by a light duty vehicle compared with . L
km- for fossil fuel-based gasoline [ ]. Most of the water used in gasoline is for the oil refining
process while most of the water used in bioethanol production is water used to grow the crop.
The water footprint includes three components green, blue, and gray WFs. Green WFs refer
to rainwater transpired and blue WFs to surface and groundwater evaporated following their
use in irrigation. Gray WF refers to water that becomes polluted during crop production and
includes the amount of water necessary to reduce pollutants through dilution discharged so
that water quality meets appropriate standards [ ].
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The global annual mean WF was Gm yr- % green, % blue, % gray and agricul‐
tural production contributed % to that total during the period - , with the remainder
accounted for by food processing and end-user consumption [ ]. The average consumer in
the United States has a WF of m yr , whereas the average consumer in China and India
-

have WFs of and m yr , respectively. The differences are mainly due to differences
-

in meat consumption. Globally, consumption of cereals gives the largest contribution to WF

of the average consumer % , followed by meat % and milk products % [ ].
“pproximately one third of the total WF of agriculture is related to livestock products. The
average WF per calorie or MJ for beef is times larger than cereals and starchy roots, and
the WF g of protein for chicken meat, eggs, and milk is . times larger than for legumes [ ].
-

The WF of any livestock product is larger than any WF of crop products of equivalent
nutritional value e.g., calories, protein, and fat due to the unfavourable feed conversion
efficiency for livestock products [ ]. The weighted global average WF of sugarcane was
m Mg- m Mg- for sugarbeet and m Mg- for maize [ ] for bioethanol. Incentives
to switch from overhead sprinkler irrigation systems to drip irrigation can potentially use %
less water and lower energy requirement for maize farmers by reducing energy needed to
pump water, and by reducing evaporation losses [ ].

. Improving the energy efficiency of food and biofuel systems

The following are some suggestions to improve the energy efficiency of food and biofuel
systems

. i Commercial hybrids of wheat and rice and ii increased crop stress tolerance. Increas‐
ing productivity during the Green Revolution was largely through a combination of crop
genetic improvement through the development of F hybrid varieties of maize and semi-
dwarf, disease-resistant varieties of wheat and rice and increased use of fertiliser and
irrigation [ ]. Unfortunately, crop yield increases are now slowing or have halted and
input costs such as fuel and fertiliser are increasing. Future increases in crop production
will have to come mainly from increased yield per hectare, higher cropping intensity
number of crops sown in the same field per year and to a lesser extent from cultivation
of new land [ , ]. Hybrids are the result of heterosis or the favourable combination of
dominant genes by crossing two genetically different parents. In general, hybrids provide
around % yield advantage over open-pollinated parents in maize, and around % over
inbred parents in wheat and rice [ ]. “lthough hybrid maize has been widely utilised,
adoption of hybrids in wheat and rice with the exception of China where O. indica hybrids
cover around % of area is low [ ]. Hence, future adoption of hybrid wheat and rice
may increase yield by around %. Currently, most of the major commercial genetically
modified GM crops are based on simple insertion of a gene for protective traits such as
insect toxin e.g., Bacillus thuringiensis, Bt and/or herbicide resistance e.g., glyphosate
tolerance . The next phase will be the development of abiotic stress tolerance genes such
as DroughtGard® drought tolerant hybrid maize released by Monsanto in . Not all
crop improvement initiatives need to be GM based on transgenic biotechnology. For
318 Photosynthesis

example, there are a number of heat tolerance breeding programs in wheat, chickpea, and
cotton using conventional breeding together with marker assisted selection [ - ].

. Reduced yield gaps. Yield gaps are the difference between realised farm yield and the
potential yield that can be achieved using available technologies and management. In
many irrigated cereal systems, actual yield tends to plateau at or around % of yield
potential while, in rainfed systems, average actual yields are no more than % of yield
potential [ ]. In many instances, however, even the % of yield potential is not
achieved, presumably due to technical, knowledge, climatic and biophysical constraints
[ ]. Reducing the yield gaps can potentially boost yields per unit of input, and hence,
energy conversion efficiency. International aid programs such as the Gates Foundation
and the Consultative Group on International “gricultural Research CGI“R are working
to help to close these yield gaps.

. Conservation agriculture. Zero or reduced tillage and retaining crop residues can
potentially reduce energy use and fuel use for farm machinery in agriculture by - %
[ ], as well as conserve soil moisture and sequester carbon in soil organic matter.
Conversion from conventional tillage to zero/reduced tillage can reduce on-farm GHG
emssions by - kg CO e ha- per season, since soil disturbance caused by tillage
increases soil organic carbon losses through decomposition accelerated oxidation and
physical erosion [ ]. Crop residues produced worldwide are estimated at Pg, equiv‐
alent to more than Pg carbon per year [ ]. With less than % of the global crop land
under conservation tillage, there is an opportunity for wider adoption of this practice to
improve energy- and water-use efficiency [ ], as well as reducing net GHG emissions.

. Legume rotations and N-fixing cereals. Replacing synthetic N fertilisers with legumes and
organic fertilisers e.g., animal manures and green manure crops can reduce the fossil
fuel combusted during fertiliser synthesis as well as reduce N O emissions. Soil microor‐
ganisms such as rhizobium N -fixers in legumes , arbuscular mycorrhizal fungi “M
fungi, which can improve plant P and Zn uptake and P solubilising fungi and bacteria
can form a symbiotic relationship with plant roots and act as biofertilisers and biopesti‐
cides [ , ]. Many bacteria can produce natural plant hormones such as ethylene,
cytokinins, auxins and gibberellins that can stimulate plant growth, increasing root
branching, or shoot development [ ]. Microorganisms can also affect gene expression
and activate plant defence mechanisms through systemic acquired resistance S“R [ ].
“ challenge going forward is to boost biological N fixation to levels that can substitute for
the synthetic fertilisers now used. Effective nitrogen-fixing wheat, rice, and/or maize
would be boons to reducing energy input to cropping systems for food, feed, and
bioenergy [ ]. Past attempts to develop such symbioses are not encouraging.

. Improving overall annual solar radiation use efficiency in annual crops. In locations
supporting only one crop per year, it may be possible to extend the photosynthetic period
via stay-green traits to maximise annual solar energy capture [ , ]. In other areas, it
may be possible to shorten the crop cycle to fit a second or third crop into the annual
rotation i.e. increase crop intensification [ ].
 Bioenergy 319
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. Perennial crops. Perennial crops have greater potential for a more sustainable production
due to their longer growing season than annuals, utilising more sunlight during the year,
and reduced farming operations [ ]. The development of perennial wheat by the Land
Institute in Kansas provides the opportunity to crop continuously without tillage and
reduce soil erosion [ ]. Most LC“ also show that perennial C biofuel crops such as
miscanthus and switchgrass and C“M biofuel crops like agave can provide positive
energy balances, supplement renewable energy demands, and mitigate GHG emissions
[ , ]. “n important question is whether perennial wheat can attain yields of present
wheats across a wide range of environments.
. “ lower-trophic-level society. Overall solar energy conversion efficiency in agriculture
would be increased if humans consumed more crops directly, rather than after their
processing through livestock. Even a modest replacement of energy-intensive meats with
less-energy-intensive grains, fruits and vegetables [ ] would be significant at the global
scale. In marine systems, gains in sustainability could come from harvesting lower trophic
level species such as algae, phytoplankton, and filter feeder organisms such as bivalves.
For example, over . Tg yr- of macroalgae seaweed is produced in China to be used as
food for humans, feed for marine animals, and industrial raw materials [ ]. “ factor to
be overcome is the global trend toward eating more energy-costly food as a component
of economic development.

. Hydrogen production and artificial photosynthesis

Hydrogen cells might be used to fuel future cars. “lthough H contains three times the energy
of petrol on a mass basis, . L of H , compressed at MPa, are needed to substitute for L
of petrol. H is also highly flammable and it is % more expensive to transport than natural
gas [ ]. There is interest in both biotic and abiotic sytems that mimic the biological production
of H gas via the breakdown of water analogous to its electrolysis, H O → H + O , which
is carried out in photosystem II using solar energy . Certain algae contain the enzymes
hydrogenase or nitrogenase a key enzyme in nitrogen fixation which can produce H from
CO or organic waste [ ]. The combination of microalgae harvesting radiation in the
waveband - nm and purple bacteria using the waveband - nm allow a more
complete utilisation of the solar energy spectrum. There are still many unresolved problems
in producing H using algae and bacteria, including how to combine microalgal and bacterial
biological processes [ ].
“rtificial photosynthesis involves mimicking natural systems using molecular photocatalytic
systems for light-driven water oxidation and H production [ ]. “rtificial photosynthesis
was only an academic activity until the development of the first practical artificial leaf by
Nocera in [ ]. The key to this breakthrough was the discovery of new, cheaper photo‐
catalysts made from nickel and cobalt that are capable of splitting water into H and O
efficiently. This artificial leaf was claimed to be potentially times more efficient in photo‐
synthesis than a natural leaf [ ]. Commercialisation of artificial photosynthesis is yet to be
proven.
320 Photosynthesis

. Conclusions

Solar radiation is the ultimate source of renewable energy for human use, and bioenergy will
continue to be a major vehicle for its use. Solar-energy conversion efficiency by even the most
productive plant communities are less than %, however, while photovoltaic cells may
approach %. “verage plant communities operate at considerably lower efficiencies, but
there are opportunities to substantially increase the average efficiency in crop systems.
Photosynthesis is now used extensively in agriculture to produce food, feed, fibre, and biofuels,
but the current biofuels bioethanol and biodiesel , mainly produced from first generation
feedstocks e.g., sucrose from sugarcane, carbohydrates from maize seeds, and lipids from
rapeseed seeds , constitute only a small fraction % of present transportation energy, and a
much smaller fraction of total human energy supply. The future second generation biofuels
will come from dedicated perennial energy crops e.g., miscanthus, switchgrass, agave,
pongamia , and in the near future, hydrogen gas may be produced from algae, bacteria, or
artificial photosynthesis to fuel hydrogen-cell powered cars.

Abbreviations and units

Abbreviation Term represented

Bt Bacillus thuringiensis

Btu British thermal unit (equivalent to 1.055 kJ)

CAM Crassulacean acid metabolism

CO2e Carbon dioxide equivalents

εc Photosynthetic conversion efficiency (J J-1 in resulting biomass)

εi Light interception efficiency (fraction of incident radiation absorbed by a plants photosynthetic

apparatus, J J-1)

EJ Exajoules (1018 J)

εp Partitioning efficiency or harvest index (J J-1).

FER Fossil energy ratio

GHG Greenhouse gas

GM Genetically modified

GPP Gross primary production

kcal Kilocalories

kcat Catalytic rate of carboxylation (reactions catalysed per second by each enzymatic site)

LCA Life cycle analysis

Mtoe Million Mg of oil equivalent

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NEV Net energy value

NPP Net primary production

NPQ Non-photochemical quenching

PAR Photosynthetically active radiation (radiation in the 400-700 nm wave band)

Pg Petagram (1 × 1015 g)

Pmol Petamole (1 × 1015 mol)

SAR Systemic acquired resistance

St Total incident solar radiation across the growing season (J m-2)

Tg Teragram (1 × 1012 g)

TL Trophic level

TW Terawatt (1 × 1012 W)

WF Water footprint

Y Biomass energy yield (J m-2 of ground)

τ Specificity for CO2 relative to O2

Author details

Daniel K. Y. Tan* and Jeffrey S. “mthor

*“ddress all correspondence to [Link]@[Link]

Faculty of “griculture and Environment, The University of Sydney, Sydney, NSW, “ustralia

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