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Energy I5 (1998) 205-210








Institute of Crop Science
Federal Agricultural Research Centre (FAL)
Bundesallee 50, 38 I I6 Braunschweig. German)
Tel: +49 531 596605: Fax. +49 53 I 596365

Careful selection of crop species. amoung other aspects, is very helpful in enhancing energy production b>
\<a! of increased biomass yields from agricultural land. A wide range of Cj and Cd plant species has been
tntroduced and investigated for their environmental and climatic impact. The results indicate already that
some perennial Cd crop species posses high yield potential, lower erosion-index, better CO* reduction rates
and need less fertilizer. water and chemicals. !:, 1998 Published by Elsevler Science Ltd. All rights reserved

Energy balance: photosynthetic pathways; biofuels

Nearly all of the energy which we use today has been collected and stored by the process of photosynthesis.
The amount of carbon fixed each year by autotrophic plants. in the form of biomass, is roughly 200 billion
rons or approximately ten times the energy equivalent used yearly in the world. Of these 200 billion tons.
800 million are used to feed the human population, i.e. 0.4%.
The fundamental process of biomass accumulation within the context of energy is based on photosynthesis.
The green plant is the only organism able to absorb solar energy with the help of the green pigment
chlorophyll. With the aid of carbon dioxide and water, solar energy is converted into the chemical energy
of organic compounds (Fig. I). In addition to C. H. and 0, plants also incorporate N and S into organic
matter by means of light reactions.
The photosynthetic process also requires other adequate nutrients for optimum functioning. and an
adequate temperature in which to take place. The efficiency of photosynthesis, expressed as the ratio
between chemical energy fixed by plants and the energy contained in light rays falling on plants. is less
than 1%. A small increase in this efficiency would have spectacular effects in view of the scale of the
process involved. However. what has been defined as photosynthesis efficiency is actually the sum of the
individual effectivenesses of a large number of complex and related processes taking place both within the
plants and in their environment.

front matter C

PII: SO960-l481(98)00157-8

1998 Published by Elsevier Science Ltd. All rights





If we summarize the solar energy which hits every hectare globally it averages out to be the amount found
at 40 latitude where every hectare receives 1.47 x lOI calories for total energy radiation (TER). If this
energy is completely converted into the chemical energq of carbohydrates we would expect a yield of more
than 2000 tons per hectare. But the photosynthetically acti\,e radiation (PAR) amounts to 43% of the total
solar radiation on the earths surface.


602-tCaHuOa'=-6CO2t 6 H20t
carbon dioxide

water aIII~~~I



glucose t


Fig. 1. The almost closed cycle of energy production from biomass

In the case of a crop yielding 10 tons of dry matter per hectare per year. located at 40 latitude, the
photosvnthetic efficiency would be 0.27%.for TER or 0.63 for PAR. As the PAR is about 43% of TER. the
upper jimit of efficiency for the absorption of solar radiation is 6.8% of the total radiation, or as a value of
15.8% with respect to PAR.
Considering the total amount of radiation incident at certain areas and keeping in mind that the calorific
content of dry matter is about 4000 kilocalories per kilogram (= 4 Kcal/g), a theoretical upper production
limit of about 250 tons of dry matter per hectare could be considered for an area having an amount of
incident radiation similar to that at 40 latitude. This production value has not been attained so far with
any crop, as the vegetation cycle and high growth and solar utilization rates cannot be achieved and
maintained continuously throughout the entire year.
From laboratory studies we know that the photosynthetic apparatus of green plants works with an
efficiency of more than 30%. Why this discrepancy? What we really measure when we determine the
harvested biomass of our fields is the difference between the efficiencies of photosynthesis and respiration.
Furthermore, we have to consider that the growing season of our main crops comprises just a fraction of
the year and that by mutual shading of the leaves, unfavorable water content of the soil, air humidity and
too low or too high temperatures the yield must remain below the theoretical limit. Besides this the solar
spectrum contains broad regions which are photosynthetically practically inactive (< 400 nm and > 700
nm). In addition, the availability of adequate resources of water and nutrients during the growth cycle needs
to be considered. The determinants that affect photosynthesis are divided into the three main groups of
environmental factors, genetic structure of genotype and population, and external inputs (Fig. 2). Their
influence on the plants photosynthesis controls the final biomass yield.




The two major pathways of photosynthesis are the C; pathway and the CJ pathway. A third. less common
pathway, is the CAM pathway. In the CJ pathway, the first product of photosynthesis is a j-carbon organic
acid (3-phosphoglyceric acid) while in the Ca the first products are 4-carbon organic acids (malate and
aspartate). In general the C3 assimilation pathway is adapted to operate at optimal rates under conditions of
low temperature (15-20C). The C3 species have at a given radiation level relatively lower rates of CO?
exchange than Cq species which are adapted to operate at optimal rates under conditions of higher
temperature (30-35C) and have comparatively higher rates of CO* exchange. Furthermore, CJ species
habe maximum rates of photosynthesis in the range of 70-100 mg COxidmVh with light saturation at 1.O1.J cal/cm2/min total radiation. while C3 species have maximum rates of photosynthesis in the range 1j-30
mg COz/dm2/h with light saturation at 0.2-0.6 cal/cmVmin.

Fig. 2. Determinants of potential biomass production (El Bassam, 1990)

Breeding and selection, through both natural and human agencies, have changed the temperature response
of photosynthesis in some CJ and Cd species. Consequently, there are Cj species (e.g. cotton and
groundnut) whose optimal temperature is in a medium to high range (25-30 C), and there are Cq species
(e.g. maize and sorghum), where, for temperate and tropical highland cultivars. the optimal temperature is
in a low to medium range (20-30C).
One additional group of species has evolved and adapted to operate under xerophytic conditions. These
species have the Crassulacean Acid Metabolism (CAM). Although the biochemistry of photosynthesis in
the CAM species has several features in common with Cd species, particularly the synthesis of 4-carbon
organic acids, CAM species have some unique features which are not observed in C3 and Cq species. These
include capturing the light energy during the day-time and fixing CO1 during the night-time. with
consequently very high water use efficiencies. There are two CAM species of agricultural importance (i.e.
pineapple and sisal).




The Cl, Cd and CAM crops can be roughly classified into five groups (Table 1). based on the differences
between crop species in their photosynthetic pathways and the response of photosynthesis to radiation and
CJ plant species tend to be more efficient in Mater utilization because they do not have to open their
stomata so much for inward diffusion of CO>. Thus transpiration is reduced. The corollary is that Ci
species are usually more efficient net photosynthesizers at lower temperatures than Cd species. Both
metabolic groups are capable of substantial climatic adaptation (Clayton and Renovize. 1986). ~4L~canthu.s
spp. and Spartina spp. are both CJ species and more efficient photosynthesizers than a wide variety of C;
grasses. This reflects the fact that there are possibilities for genetic selection of Cd plants with improved
cold tolerance. The Cd species may utilize nitrogen with greater efficiency.

Table I. Some physiological characteristics according to the photosynthesis-type

(FAO, 1980; revised El Bassam. 1995).
Crop yroup
- Radiation






0.2 - 0.6

0.3 - 0.8

I .o - I .4

20 - 30

40 - 50




0.6 - 1.4


- Max.

net rate of
CO? exchange at
light saturation
(mg dm h)


70 - 100


- 50

- Temperature
respon. of photosyn
optimum temp.
operative temp.
- :trq$Y


- Water use

- Crop species

15 - 20C
5 - 30C

25 - 30C
IO - 35OC

30 - 35C
15 -45C

20 - 30C
IO - 35T

25 - 35C

20 - 30 g

30 - 40 g

30 - 60 g

40 - 60 g

20 - 30 g

400 - 800




50 - 200

field mustard,
potato, oat.
flax, rape,
rye, wheat,
olive. barley,
lentil, linseed,
broom. poplar,

- 700

millet, maize.
groundnut. rice.
soybean. castor,
barnyard millet.
sweet sorghum.
sesame, tobacco,
grain sorghum.
fiber sorghum,
willow, bananas, maize, sugarcane.
aieman grass
Miscanthus spp.,
sunflower. kenaf.
Spartina spp.
sweet potato.
bamboo. cotton.
coconuI. cassava.
Eucalyprus spp.,


WREC 1998


An estimate of the upper threshold for converting incident solar energy into biomass production for C; and
Cd species was made by Hall e/ al. (1993).
For C~plants:

100% x 0.50 x 0.80 x 0.28 x 0.60 = 6.7%

0.50 = The 50% of light that is photosynthetically

between 400 and 700 nm.

0 80 = The about 80% of PAR captured by photosynthetically active compounds; the

rest 1s reflected. transmitted, and absorbed by non-photosynthesizing leaves.

0.28 = The theoretical maximum

absorbed PAR to glucose. 28%.


active radiation (PAR). wavelengths

energy efficiency

of converting

the effectively

0.60 = The about 60% energy stored in photosynthesis remaining after 40% is
consumed during dark respiration to sustain plant metabolic processes.

The 6.7% applies to Ca plants (plants where the first stable products of photosynthesis

are 4-carbon

FOI- Cj plants. 2 additional losses occur: Cj plants lose about 30% of the already fixed CO2 during
photorespiration. and Cj plants become light-saturated at lower light intensities than Cd plants. so that Cj
plants are unable to utilize perhaps 30% of the light absorbed by photosynthetically active compounds.
For Cl plants: 6.7% x 0.70 x 0.70 = 3.3%
These figures can be now used to calculate the potential dry matter yield for each biomass crop. However.
the following information is also required:

Length of growing season.

Average daily solar radiation in the region,
Energy content of the feedstocks,
Partitioning of the biomass above and below ground. and
Availability of water and nutrients.

The final biomass accumulation is then determined by the genetic structure of the genotype and population.
Plants with the Cd photosynthetic pathway have a maximum possible potential in temperate climates of 55
tonnes per hectare per year as compared to 33 tonnes per hectare per year for temperate C, crops (Bassham.
1980). The potentials could be much higher for warmer and more humid climates. Samson and Chen
(1995) estimated the maximum photosynthetic efficiency of the biomass plantation studied in Canada
would be 64.8 ODTlha for switchgrass (Cd) and 33.6 ODT/ha for Willow (Cj). ODT being oven dried tonne.

1he growth and productivity of crops depends on their genetic potential. The degree of the realization of
this potential is closely related to the environmental factors dominating in the region and the external
inputs. Yields achieved in field experiments do not represent the physiological limits of the present
cultivars, but only demonstrate that portion of the genetic potential which is realized by optimal utilization
of present means of cultivation and levels of inputs (Dambroth and El Bassam, 1990). Yield and response
of cultivars to environmental conditions and inputs is under genetic control and therefore an improved
response is accessible via screening, selection and breeding.
Climate and local soil properties are the most important constraints influencing the biomass productivity
within a specific region. Temperature, precipitation and global radiation are the few classical climatic
factors which influence, to a great extent, the regional distribution of various plant species. Breeding and
selection. through both natural and human activities, has changed the temperature response of
photosynthesis in some Cs and Cd species. Consequently, there are C3 species whose optimum temperature
is in a medium to high range (25-30C). and there are Cd species (e.g. maize. sorghum) where. for
temperate and tropical highland cuhivars, the optimum temperature is in the range (20-30C).

WREC 1998


The criteria for an ideal fuel crop can be summarized as:


Efficient conversion of solar radiation to biomass. with a yield close to the theoretical maximum. 55 t
DM/ha for C4 crops and 35 t DM/ha for C3 crops. or crops with high oil contents or high fermentable
constituents (sugar, starch, etc.).
High dry matter content (%).
Efficient use of minerals and water.
Convenient energy balance (output. input).
IIigh energy density (MJ/kg).
Enable sustainable production systems.
Good disease resistance.
Consistent to the environment and climate.

[f all these aspects would be considered, the best fuel crop would be characterized as a high productive
perennial plant species. They can be high productive perennial Cj plant species such as Arundo donux or
CAplant species such as Miscanrhus sinensis.
The reasons are:

Relatively low water and fertilizer requirements

Low herbicide requirements
No need for fungicide, insecticide and nematodicide
Energy balance is very suitable (positive)
Very low erosion risk
Landscape: pleasant visual impact - possible integration with other species
Habitat for birds, mammals and other organisms
Total crop processing possible
Sustainable production system: conservation of resources and environmentally consistent
Positive impact on climate

These crops are basically lignocellulosic plant species. The biomass can be converted to a wide variety of

Bassham, J.A. (1980) Energy crops (energy farming). In: Biochemical und Phofosynfhefic Aspects of
Energy Producfion 6. Academic Press, New York. pp. 147-173.
Clayton. W.D. and S.A. Renovize (1986) Genera Graminum, Grasses of fhe World. Kew Bulletin
Additional Series XIII.
Dambroth, M. and N. El Bassam (1990) Genotypic variation in plant productivity and consequences for
breeding of low-input cultivars. Generic Aspecfs of Plant Minerul Nufrifion, Developmenfs in
Plant and Soil Sciences, Kluwer Academic Publishers. pp. 1-7.
El Bassam, N. (1990) Requirements for biomass production with higher plants under artificial ecosystems.
Workshop Artificial ecological systems. Marseille. pp. 59-65.
El Bassam, N. (1995) Auswirkungen einer Klimaverlnderung auf den Anbau von C4-Pflanzenarten zur
Erzeugung von Energietrlgem. Miff. Ges. Pfunzenbuuwiss. 8: 169- 172.
FAO (1980) Report on the agro-ecological zones project, Vol. 1. FAO, Rome. pp. 24-48.
Hall. D.O.. F. Rosillo-Calle, R.W. Williams and J. Woods (1993) Biomass for energy: supply prospects.
In: T.B. Johannsson et al., (eds). Renewable Energy: Sources of Fuels and Elecfricify. Island Press,
Washington, D.C. pp. 593-651.
Samson, R. and Y. Chen (1995) Short-rotation forestry and water problem. Canadian Energy Plantation
Workshop. pp. 43-49.