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Applied Ecology (BOT-7203)

Assignment topic:
“Physiological and ecological considerations: Light leaves and
photosynthesis, photosynthetic response to light by intact
leaves, CO2 climatic changes in CO2 and photosynthetic
response of C3, C4 and CAM plants”

Submitted by:
Ansa Yousaf

Roll no: 0361MPHIL-BOT-23

MPhil Botany (evening)

1st Semester

Submitted to:
Sir Tariq Rizwan

Department of Botany, Government College University, Lahore

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Table of contents
Photosynthesis: Physiological and Ecological Considerations

PHOTOSYNTHESIS: PHYSIOLOGICAL AND ECOLOGICAL CONSIDERATIONS

Photosynthesis is a complex process that converts solar energy to organic compounds' chemical energy.
It occurs in organisms that continuously respond to internal and external changes. The impact of the
environment on photosynthesis is important for plant physiologists and agronomists, as plant productivity
depends on prevailing photosynthetic rates in a dynamic environment. British plant physiologist F.F.
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Blackman's 1905 hypothesis suggests that photosynthesis can be limited by light or CO2 concentration,
but not both factors.

In the intact leaf, three major metabolic steps have been identified as important for optimal photosynthetic
performance:

1. Rubisco activity

2. Regeneration of ribulose bisphosphate (RuBP)

3. Metabolism of the triose phosphates

The first two steps are the most prevalent under natural conditions. Table provides some examples of
how light and CO2 can affect these key metabolic steps. In the following sections, biophysical,
biochemical, and environmental aspects of photosynthesis in leaves are discussed in detail.

LIGHT, LEAVES, AND PHOTOSYNTHESIS

The leaf adds new levels of complexity to photosynthesis. At the same time, the structural and functional
properties of the leaf make possible other levels of regulation. We will start by examining how leaf
anatomy, and movements by chloroplasts and leaves, control the absorption of light for photosynthesis.
Then we will describe how chloroplasts and leaves adapt to their light environment and how the
photosynthetic response of leaves grown under low light reflects their adaptation to a low-light
environment. Leaves also adapt to high light conditions, illustrating that plants are physiologically
flexible and that they adapt to their immediate environment.
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Both the amount of light and the amount of CO2 determine the photosynthetic response of leaves. In
some situations, photosynthesis is limited by an inadequate supply of light or CO2. In other situations,
absorption of too much light can cause severe problems, and special mechanisms protect the
photosynthetic system from excessive light. Multiple levels of control over photosynthesis allow plants
to grow successfully in a constantly changing environment and different habitats.

CONCEPTS AND UNITS IN THE MEASUREMENT OF LIGHT

Light parameters are crucial in measuring light, including spectral quality, amount, and direction. Flat or
planar light sensors are best suited for flat leaves, as they can measure the amount of energy that falls on
a flat sensor per unit time. Units can be expressed in terms of energy, such as watts per square meter (W
m–2) or moles per square meter per second (mol m–2 s–1).

Light can also be measured as the number of incident quantum (singular quantum), expressed in moles
per square meter per second (mol m–2 s–1). This measure is called photon irradiance. Quanta and energy
units can be inter converted relatively easily, provided that the wavelength of the light, l, is known. The
energy of a photon is related to its wavelength, with c being the speed of light, h being Planck’s constant,
and l being the wavelength.
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The direction of light can be affected by the light intercepting object being not flat, such as complex
shoots, whole plants, or chloroplasts. In some situations, light can come from many directions
simultaneously, making it more sense to measure light with a spherical sensor that takes measurements
omnidirectionally. The term for this omnidirectional measurement is fluence rate, which can be expressed
in watts per square meter (W m–2) or moles per square meter per second (mol m–2 s–1).

In contrast to a flat sensor's sensitivity, the sensitivity to light of a spherical sensor is independent of
direction. The sensitivity to light depends on whether the light is collimated (rays are parallel) or not.

• Leaf Anatomy Maximizes Light Absorption

About 1.3 kW m–2 of radiant energy from the sun reaches Earth, but only about 5% can be converted
into carbohydrates by a photosynthesizing leaf. This is because a significant fraction of the incident light
is either too short or too long to be absorbed by the photosynthetic pigments. A significant fraction is lost
as heat and a smaller amount is lost as fluorescence. Only photons of wavelengths from 400 to 700 nm
are utilized in photosynthesis, with about 85 to 90% of this PAR absorbed by the leaf. The transmitted
and reflected light are vastly enriched in green, resulting in the green color of vegetation.
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The leaf's anatomy is highly specialized for light absorption, with the
outermost cell layer, the epidermis, typically transparent to visible
light. Convex epidermal cells can act as lenses, focusing light so that
the amount reaching some chloroplasts can be many times greater
than the amount of ambient light. This is common among herbaceous
plants and particularly prominent among tropical plants growing in
forest understory, where light levels are very low.

Below the epidermis, the top layers of photosynthetic cells are called
palisade cells, shaped like pillars standing in parallel columns one to
three layers deep. Some leaves have several layers of columnar
palisade cells, allowing more light than expected to penetrate the first layer of palisade cells due to the
sieve effect and light channeling.

The sieve effect occurs when chlorophyll is not uniformly distributed throughout cells but is confined to
chloroplasts, creating gaps where light is not absorbed. This results in less total absorption of light by a
given amount of chlorophyll in a palisade cell than by the same amount in a solution. Light channeling
occurs when incident light is propagated through the central vacuole of the palisade cells and through air
spaces between the cells, facilitating light transmission into the leaf interior.

Below the palisade layers is the spongy mesophyll, where cells are irregular in shape and surrounded by
large air spaces. These large air spaces generate many interfaces between air and water that reflect and
refract light, causing light scattering. This phenomenon is especially important in leaves, as it increases
the length of the path over which photons travel, increasing the probability of absorption. The palisade
cell properties that allow light to pass through and the spongy mesophyll cell properties that facilitate
light scattering result in more uniform light absorption throughout the leaf.
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Deserts have high light levels that can harm leaves. To mitigate this, leaves have special anatomic features
like hairs, salt glands, and epicuticular wax that increase light reflection, reducing absorption by up to
40%. This minimizes heating and other issues associated with excessive light absorption.

• Chloroplast Movement and Leaf Movement Can Control Light Absorption

Chloroplast movement is a common phenomenon in higher plants, including algae, mosses, and leaves.
It allows plants to regulate light absorption by controlling their orientation and location. Under low light,
chloroplasts gather at cell surfaces parallel to the leaf plane, maximizing absorption. Under high light,
they move to cell surfaces parallel to the incident light, reducing light absorption by about 15%.
Chloroplast movement is a blue-light response, with some algae controlling it through phytochrome. In
leaves, chloroplasts move along actin microfilaments in the cytoplasm, and calcium regulates their
movement.

Leaves have the highest light absorption when the leafblade or lamina is perpendicular to the incident
light. Some plants, such as Alfalfa, cotton, soybean, bean, lupine, and some wild species of the mallow
family (Malvaceae), can control light absorption through solar tracking. Solar-tracking leaves maintain
a nearly vertical position at sunrise, facing the eastern horizon, where the sun will rise. They track the
sun only on clear days and stop when a cloud obscures the sun.

Solar tracking is another blue-light response, with the sensing of blue light occurring in specialized
regions. Some plants can also move their leaves to avoid full sunlight exposure, minimizing heating and
water loss. These leaves are called paraheliotropic, while those that maximize light interception by solar
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tracking are called diaheliotropic. Solar tracking gives plants an advantage in arid regions, as it maintains
maximum photosynthetic rates throughout the day and lowers air temperature during early morning and
late afternoon.

• Plants Adapt to Sun and Shade

Plants have the ability to adapt to various light regimes, growing as sun plants in sunny areas and shade
plants in shady habitats. Some shade plants receive less than 1% of the PAR available in an exposed
habitat. Sun and shade leaves have contrasting characteristics, such as more total chlorophyll per reaction
center, higher ratio of chlorophyll b to chlorophyll a, and thinner leaves. Sun leaves have more rubisco
and a larger pool of xanthophyll cycle components.

Contrasting anatomic characteristics can also be found in leaves of the same plant exposed to different
light regimes. Sun-grown leaves are thicker and have longer palisade cells, while shade-grown leaves
have cells in the upper surface with characteristics of leaves grown in full sunlight. These morphological
and biochemical modifications are associated with specific functions, such as maintaining a better
balance of energy flow through the two photosystems. Some shade plants show a 3:1 ratio of photosystem
II to photosystem I reaction centers, while others add more antennae chlorophyll to PSII.
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• Plants Compete for Sunlight

Plants compete for sunlight, with leaves forming a canopy that absorbs light and influences
photosynthetic rates and growth. Some plants have thick leaves that transmit little light, while others,
like the dandelion, have a rosette growth habit.

Trees are an excellent adaptation for light interception, with their elaborate branching structure
increasing sunlight absorption. Sunflecks, patches of sunlight that pass through small gaps in leaf canopy,
can change photon flux on leaves in dense forests more than tenfold within seconds. Sunflecks also play
a role in carbon metabolism in lower leaves in dense crops. Plant physiologists and ecologists are
interested in the rapid responses of photosynthetic apparatus and stomata to sunflecks, as they represent
unique physiological responses for capturing a short burst of sunlight.

PHOTOSYNTHETIC RESPONSES TO LIGHT BY THE INTACT LEAF

This section discusses the photosynthetic properties of plants, highlighting their adaptations to light. It
discusses typical responses to light, the light compensation point, and the quantum yields of
photosynthesis in intact leaves. It also discusses the differences in yields between C3 and C4 plants, leaf
adaptations to excess light, and different pathways of heat dissipation in the leaf.
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• Light-Response Curves Reveal Photosynthetic Properties

Photosynthesis is a crucial process in plants, where the amount of light received during the growing
season influences the photosynthetic properties of leaves. The light-response curves provide valuable
information about the photosynthetic properties of leaves, with the light compensation point being the
point at which CO2 uptake balances release. The photon flux at which different leaves reach this point
varies with species and developmental conditions.

Sun plants have higher light compensation points than shade plants, as their respiration rates are very
low, allowing them to survive in light-limited environments. As photon flux increases above the light
compensation point, it results in a proportional increase in photosynthetic rate, leading to a linear
relationship between photon flux and photosynthetic rate.

The maximum quantum yield of photosynthesis for a leaf varies between 0.04 and 0.06. In intact leaves,
measured quantum yields for CO2 fixation range between 0.04 and 0.06. Quantum yields vary with
temperature and CO2 concentration due to their effect on the ratio of carboxylase and oxygenase
reactions of rubisco.

At higher photon fluxes, the photosynthetic response to light starts to level off and reaches saturation.
Once the saturation point is reached, further increases in photon flux no longer affect photosynthetic
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rates, indicating that factors other than incident light, such as electron transport rate, rubisco activity, or
the metabolism of triose phosphates, have become limiting to photosynthesis.

Shade plants have substantially lower light saturation levels than sun plants, reflecting the maximal
photon flux to which the leaf was exposed during growth. The light-response curve of most leaves
saturates between 500 and 1000 µmol m–2 s–1, while photon fluxes well below full sunlight (about 2000
µmol m–2 s–1) are observed.

Crop productivity is related to the total amount of light received during the growing season, and given
enough water and nutrients, the more light a crop receives, the higher the biomass.

• Leaves Must Dissipate Excess Light Energy

Excess light in plants can cause damage to the photosynthetic apparatus. The xanthophyll cycle, which
includes carotenoids violaxanthin, antheraxanthin, and zeaxanthin, is an important pathway for
dissipating excess light energy. Under high light, violaxanthin is converted to antheraxanthin and then to
zeaxanthin. Zeaxanthin is the most effective in heat dissipation, with levels increasing at high irradiances
and decreasing at low irradiances. In leaves growing under full sunlight, zeaxanthin and antheraxanthin
can make up 60% of the total xanthophyll cycle pool at midday. This allows a substantial amount of
excess light energy to be dissipated as heat, preventing damage to the chloroplast's photosynthesis. The
fraction of light energy dissipated depends on factors such as irradiance, species, growth conditions,
nutrient status, and ambient temperature.

The xanthophyll cycle also operates in plants that grow in low light, such as conifers, which maintain
green leaves during winter. Zeaxanthin remains high all day, maximizing dissipation of light energy and
protecting the leaves against photooxidation. In addition to protecting the photosynthetic system against
high light, the xanthophyll cycle may also help protect against high temperatures, as chloroplasts become
more tolerant of heat when they accumulate zeaxanthin.
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PHOTOSYNTHETIC RESPONSES TO CARBON DIOXIDE

Plant growth and leaf anatomy are influenced by light, and CO2 concentration affects photosynthesis.
CO2 diffuses from the atmosphere into leaves through stomata, intercellular air spaces, and cells. High
CO2 concentrations support higher photosynthetic rates, while low concentrations limit it. This section
discusses atmospheric CO2 concentration, its availability for carbon-fixing processes, and the limitations
of CO2 on photosynthesis and the impact of CO2-concentrating mechanisms in C4 plants.

• Atmospheric CO2 Concentration Keeps Rising

Carbon dioxide is a trace gas in the atmosphere, presently accounting for about 0.037%, or 370 parts per
million (ppm), of air. The partial pressure of ambient CO2 (Ca) varies with atmospheric pressure and is
approximately 36 pascals (Pa) at sea level. Water vapor usually accounts for up to 2% of the atmosphere
and O2 for about 20%. The bulk of the atmosphere, nearly 80%, is nitrogen. The current atmospheric
concentration of CO2 is almost twice the concentration that has prevailed during most of the last 160,00
years, as measured from air bubbles trapped in glacial ice in Antarctica

Except for the last 200 years, CO2 concentrations during the recent geological past have been low,
fluctuating between 180 and 260 ppm. These low concentrations were typical of times extending back to
the Cretaceous, when Earth was much warmer and the CO2 concentration may have been as high as 1200
to 2800 ppm (Ehleringer et al. 1991). The current CO2 concentration of the atmosphere is increasing by
about 1 ppm each year, primarily because of the burning of fossil fuels. Since 1958, when systematic
measurements of CO2 began at Mauna Loa, Hawaii, atmospheric CO2 concentrations have increased by
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more than 17% (Keeling et al. 1995), and by 2020 the atmospheric CO2 concentration could reach 600
ppm.

• The greenhouse effect

The greenhouse effect, a phenomenon resulting from the trapping of long-wavelength radiation by the
atmosphere, is causing significant global warming. A greenhouse roof transmits visible light, which is
absorbed by plants and other surfaces, and converted into heat. Glass, due to its poor transmission, cannot
leave the greenhouse, leading to increased heat. The greenhouse effect also affects photosynthesis, with
CO2 and methane playing similar roles. At current atmospheric CO2 concentrations, photosynthesis in
C3 plants is limited, but as CO2 concentrations increase, this situation could change. Under laboratory
conditions, most C3 plants grow 30 to 60% faster when CO2 concentration is doubled (to 600-700 ppm),
depending on nutrient status. However, this enhanced growth is temporary. For crops like tomatoes,
lettuce, cucumbers, and roses growing in greenhouses under optimal nutrition, carbon dioxide enrichment
in the greenhouse environment results in increased productivity. The photosynthetic performance of C3
plants under elevated CO2 is enhanced due to decreased photorespiration.

CO2 Diffusion and Limitations

• CO2 is essential for photosynthesis, and its diffusion into leaves is a critical step in the process

• The primary entry point for CO2 is through stomata, followed by diffusion through intercellular
air spaces and ultimately into cells and chloroplasts

• The diffusion pathway of CO2 involves several points of resistance, including the boundary layer,
stomata, intercellular air spaces, and the liquid phase within the leaf(Evans et al., 2009)
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CO2 Limitation and Photosynthetic Performance

• The concept of the CO2 compensation point, where photosynthesis and respiration balance at a
specific CO2 concentration, illustrates the influence of CO2 availability on plant metabolism

• Understanding the dynamics of CO2 diffusion and its limitations provides valuable insights into
how changes in atmospheric CO2 levels may impact plant performance and ecosystems in the
future

• Ongoing research aims to unravel the complexities of this relationship and assess the broader
implications for global climate and agriculture(Evans et al., 2009)

Photosynthetic Response of C3, C4, and CAM Plants

• C3 Plants

Photosynthesis in C3 plants is stimulated by increasing atmospheric CO2 above the compensation point,
which is limited by the carboxylation capacity of rubisco at low to intermediate CO2 concentrations. At
high CO2 concentrations, photosynthesis becomes limited by the Calvin-Benson cycle's capacity to
regenerate ribulose 1,5-bisphosphate, which depends on electron transport rates. However,
photosynthesis continues to increase with increasing CO2 because carboxylation replaces oxygenation
on rubisco. Most leaves regulate their internal partial pressure for CO2 (ci) to maintain an intermediate
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concentration between the limits imposed by carboxylation capacity and the capacity to regenerate
ribulose 1,5-bisphosphate.

A plot of CO2 assimilation as a function intercellular partial pressures of CO2 reveals interesting
differences between C3 and C4 plants. In C3 plants, photosynthetic rates saturate at ci values of about 15
Pa, reflecting effective CO2-concentrating mechanisms. In C3 plants, increasing ci levels continue to
stimulate photosynthesis over a much broader CO2 range. The CO2 compensation point in C4 plants is
zero or nearly zero, reflecting their very low levels of photorespiration.

C3 plants may benefit more from ongoing increases in atmospheric CO2 concentrations, while C4 plants
do not benefit much due to their CO2-saturated at low concentrations. C3 photosynthesis is typically
CO2-diffusion limited today, but C3 plants still account for nearly 70% of the world's primary
productivity. The evolution of C4 photosynthesis is one biochemical adaptation to a CO2-limited
atmosphere, possibly 10 to 15 million years ago.

• C4 Plants

C4 photosynthesis, a major photosynthetic pathway, first emerged on Earth's ancient Earth over 50
million years ago when atmospheric CO2 concentrations were above current levels. C4 plants were most
favored in warm regions with the highest temperatures, such as subtropical grasslands and savannas. C4
photosynthesis was more prominent during glacial periods when atmospheric CO2 levels were below
200 ppm than it is today.

C4 plants' CO2-concentrating mechanisms allow them to maintain high photosynthetic rates at lower ci
values, requiring lower stomatal conductance for photosynthesis. This allows C4 plants to use water and
nitrogen more efficiently than C3 plants. However, the additional energy cost of the concentrating
mechanism makes C4 plants less efficient in their light utilization, which is likely one reason why most
shade-adapted plants in temperate regions are C3 plants.
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In summary, C4 photosynthesis became a major photosynthetic pathway on Earth's ancient Earth when
global CO2 concentrations decreased below a critical threshold. The negative impacts of high
photorespiration and CO2 limitation on C3 photosynthesis were greatest under warm to hot growing
conditions, especially when atmospheric CO2 was reduced.

• CAM plants

Plants with crassulacean acid metabolism (CAM), including many cacti, orchids, bromeliads, and other
succulents, have stomatal activity patterns that contrast with those found in C3 and C4 plants. CAM
plants open their stomata at night and close them during the day, exactly the opposite of the pattern
observed in leaves of C3 and C4 plants. At night, atmospheric CO2 diffuses into CAM plants where it is
combined with phosphoenolpyruvate and fixed into malate . The ratio of water loss to CO2 uptake is
much lower in CAM plants than it is in either C3 or C4 plants. This is because stomata are primarily open
only at night, when lower temperatures and higher humidity contribute to a lower transpiration rate.

The main photosynthetic constraint on CAM metabolism is that the capacity to store malic acid is limited,
and this limitation restricts the total amount of CO2 uptake. However, some CAM plants are able to
enhance total photosynthesis during wet conditions by fixing CO2 via the Calvin–Benson cycle at the
end of the day, when temperature gradients are less extreme. In water-limited conditions, stomata open
only at night.

Cladodes (flattened stems) of cacti can survive after detachment from the plant for several months
without water. Their stomata are closed all the time, and the CO2 released by respiration is refixed into
malate. This process, which has been called CAM idling, also allows the intact plant to survive for
prolonged drought periods while losing remarkably little water.

Evolutionary Considerations
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• C3 photosynthesis is the ancestral pathway, with C4 photosynthesis evolving more recently, about
10 to 15 million years ago

• Understanding the photosynthetic responses of C3, C4, and CAM plants provides valuable
insights into their ecological niches, resource utilization, and adaptations to varying
environmental conditions (Bowling et al., 2008).

FACE Experiment: Investigating the Impact of Elevated CO2 on Photosynthesis and Respiration

• The Free-Air CO2 Enrichment experiments represent a groundbreaking initiative aimed at

unraveling the intricate interactions between plant physiology and atmospheric CO 2
concentrations

• As we look toward the future with CO2 levels surpassing 400 ppm, the FACE experiments provide
valuable insights into the responses of different plant types (Long et al., 2004).

Responsive Nature of C3 and C4 Plants

• Early observations from FACE experiments reveal distinct responses between C3 and C4 plants

• However, an intriguing down-regulation of photosynthetic capacity is evident, showcasing

reduced enzyme activity associated with the dark reactions of photosynthesis

• The nuanced reactions of both C3 and C4 plants shed light on the intricate dynamics of
photosynthetic acclimation to changing atmospheric CO2

Influencing Factors Beyond CO2

• While CO2 is a pivotal factor, FACE observations underscore the multifaceted nature of plant
growth responses

• Nutrient availability quickly emerges as a limiting factor, challenging the initial expectations set
by greenhouse studies
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• The interplay of these factors gains significance in the context of rising temperatures and changing
climate conditions associated with elevated CO2 levels

Respiration Dynamics and Acclimation

• Elevated CO2 levels not only impact photosynthesis but also induce changes in plant respiration

• This rise in temperature, in turn, influences basic mitochondrial respiration and the respiration
rates of soil microbes and fungi

• While respiration rates are altered compared to today's atmospheric conditions, they do not surge
as predicted without the down-regulation acclimation response
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Summary

Photosynthesis is the primary function of leaves, which absorb light energy and convert about 5% of
solar energy into carbohydrates. In dense forests, almost all PAR is absorbed by leaves, and within a
canopy, leaves maximize light absorption through solar tracking and chloroplast movements. Some plants
respond to various light regimes, with sun and shade leaves having contrasting biochemical
characteristics.

Light response curves show irradiance where photosynthesis is limited by light or CO2. Shade plants
have lower light compensation points due to low respiration rates. The quantum yield of C3 plants is
higher below 30°C than that of C4 plants, while above 30°C, the situation reverses. Factors beyond the
saturation point limit photosynthesis, such as electron transport, rubisco activity, or triose metabolism.

Plants are plastic in their adaptations to temperature, with optimal photosynthetic temperatures having
strong genetic and environmental components. Leaf absorption generates a heat load that must be
dissipated. Temperature sensitivity curves identify a range for optimal photosynthesis, optimal
photosynthesis, and destructive events.

Photosynthetic responses to carbon dioxide have been increasing since the Industrial Revolution due to
human use of fossil fuels. Concentration gradients drive the diffusion of CO2 from the atmosphere to
rubisco, effectively providing energy for photosynthesis. C3 plants are more responsive to elevated CO2
than C4 plants.

The carbon isotope ratios of leaves can be used to distinguish photosynthetic pathway differences among
different plant species. Both C3 and C4 plants have less 13C than does CO2 in the atmosphere, indicating
that leaf tissues discriminate against 13C during photosynthesis.
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Reference

• Ainsworth, E. A., and Rogers, A. (2007) The response of photosynthesis and stomatal
conductance to rising [CO2]. Plant Cell Environ. 30: 258–270.

• Adams, W. W., Demmig-Adams, B., Rosenstiel, T. N., and Ebbert, V. (2001) Dependence of
photosynthesis and energy dissipation activity upon growth form and light environment during
the winter. Photosyn. Res. 67: 51–62.

• Long, S. P., Ainsworth, E. A., Leakey, A. D., Nosberger, J., and Ort, D. R. (2006) Food for thought:
Lowerthan-expected crop stimulation with rising CO 2 concentrations. Science 312: 1918–1921.

• Bowling, D. R., Pataki, D. E., and Randerson, J. T. (2008) Carbon isotopes in terrestrial ecosystem
pools and CO2 fluxes. New Phytol. 178: 24–40.

• von Braun, S. S., and Schleiff, E. (2007) Movement of endosymbiotic organelles. Curr. Protein
Pept. Sci. 8: 426–438.

• Keeling, C. D., Piper, S. C., Bacastow, R. B., Wahlen, M., Whorf, T. P., Heiman, M., and Meijer,
H. A. (2005) Atmospheric CO2 and 13 CO2 exchange with the terrestrial biosphere and oceans
from 1978 to 2000: Observations and carbon cycle implications. In A History of Atmospheric
CO2 and its Effects on Plants, Animals, and Ecosystems, J. R. Ehleringer, T. E. Cerling, and M.
D. Dearing, eds., Springer-Verlag, New York, pp 83–112.

• Vogelmann, T. C., and Han, T. (2000) Measurement of gradients of absorbed light in spinach
leaves from chlorophyll fluorescence profiles. Plant Cell Environ. 23: 1303–1311.

• von Caemmerer, S. (2000) Biochemical Models of Leaf Photosynthesis. CSIRO, Melbourne,

Australia.

• Evans, J. R., Kaldenhoff, R., Gentrry, B., and Terashima, I. (2009) Resistances along the CO 2
diffusion pathway inside leaves. J. Exp. Bot. 60: 2235–2248.
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