UNIT II

Biochemistry and Metabolism

Chapter 9: Photosynthesis:
Physiological and Ecological
Considerations

Scanning electron micrographs of the leaf anatomy of a legume

acclimation

Shade leaf

Sun leaf

Leaf movement in sun-tracking plants

Solar tracking in many plants: cotton, soybean, bean, lupine, alfalfa

Leaf angle and leaf movement can control light absorption

Leaf movement

Heliotropism: bending toward the sun;

Diaheliotropic: leaves that maximize light interception
by solar tracking;
Paraheliotropic: leaves move to avoid full exposure to
sunlight, minimizing heating and water loss.
Acclimation: a growth process in which each newly
produced leaf has a set of biochemical and
morphological characteristics suited to the particular
environment in which it unfolds.
Shade leaves: more total chlorophyll per reaction
center, a higher ratio of chlorophyll b to a; thinner;
Sun leaves: more rubisco, a larger pool of xanthophyll
cycle components than shade leaves

Chloroplast distribution in photosynthesizing cells of the duckweed Lemna

In (A) and (B), chloroplasts are positioned near the upper surface of the cells to
absorb maximum amounts of light;
(C) The chloroplasts move to the side walls, where they shade each other to
minimize the absorption of excess light.

Response of photosynthesis to light in a C3 plant

Changes in photosynthesis as a function of intercellular CO 2 concentrations

In C4 plants, photosynthetic rates saturate at ci ~15 Pa due to its CO2 concentrating

mechanism; the CO2 compensation point is zero or near zero, very low level of
photorespiration. In C3 plants, increasing c i levels stimulate photosynthesis; the
CO2 compensation point is ~ 10 Pa.

Free Air CO2 Enrichment (FACE) experiments

FACE experiments
showed that C3 plants
are more responsive to
elevated [CO2] than are
C4 plants

Chapter 9

Photosynthesis is the primary function of leaves

Irradiance: amount of energy that fall on a flat sensor of
known area per unit time, unit: W/m2 (1W = 1 joule[J] s-1).
Photon irradiance: number of incident quanta striking the
leaf. Unit: mol m-2 s-1
(1 mole of light = 6.02 * 1023 photons)
Energy of a photon:
E = hc/
h: Plancks constant (6.63 * 10-34);
c: speed of light (3 * 108 m s-1);
: the wavelength of light (1 nm = 10-9 m);
Photosynthetically active radiation (PAR): 400 700 nm.
Commonly expressed as mol m-2 s-1

Incident sunlight relative to leaf angle

Photosynthesis

Limiting factor

Light
Temperature
[CO2]
Water

Rubisco activity
Regeneration of RuBP
Metabolism of the triose phosphates
Under any particular conditions, the rate of
photosynthesis is limited by the slowest step in the
process limiting factor
Supply / Demand

Optical properties of a bean leaf

50% of light can not be absorbed by leaf;

15% of PAR is reflected or transmitted;
> 50% PAR absorbed is lost as heat or florescence;
<5% of the incident energy is converted into the
energy stored as carbohydrate

Leaf anatomy
Palisade cells; spongy mesophyll cells
Sieve effect: uneven distribution of chlorophyll
causes light transmission
Light channeling: incident light is propagated through
the central vacuoles of the palisade cells, the air
spaces between cells, an arrangement that facilitates
the transmission of light into the leaf interior.
Interface light scattering: light reflected or refracted in
the interfaces between air and water in the spongy
mesophyll tissues

The spectral distribution of sunlight at the top of a canopy and under the canopy

For unfiltered sunlight, the total irradiance was 1900 - 2000 mol m-2 s-1
(~900 W m-2); for shade: 17.7 mol m-2 s-1.

Light-response curves of photosynthetic carbon fixation in sun and shade plants

Typically, shade
plants have low light
compensation points
and have lower
maximal
photosynthetic rates
than sun plants

The quantum yield of photosynthetic carbon fixation in a C3 plant and a C4 plant

Photorespiration increases with temperature in C3 plants

and the energy cost of net CO2 fixation increases.
At lower temperature C3 plant photosynthesis is more
efficient

Light-response of photosynthesis of a sun plant grown under sun or shade conditions

Beyond the saturation

point, factors other than
incident light, such as
electron transport,
rubisco activity, or triose
metabolism, limit
photosynthesis

Changes in photosynthesis in Sitka spruce as a function of irradiance

In a canopy, as a result of shading, much higher

irradiance levels are needed to saturate photosynthesis
Photosynthesis of the intact plant is the sum of the
photosynthetic activity of all the leaves

Excess light energy in relation to a light-response curve

Diurnal changes in xanthophyll content as a function of irradiance in sunflower

Changes over the course of a day in allocation of photons absorbed by sunlight

Dynamic photoinhibition:
temporarily diverts excess
light absorption to heat but
maintains maximal
photosynthetic rate

Chronic photoinhibition:
exposure to high levels of
excess light that damage the
photosynthetic systems and
decrease both quantum
efficiency and maximum
photosynthetic rate

The absorption and dissipation of energy from sunlight by the leaf

Radiative heat loss

Sensible heat loss

Latent heat loss

Bowen ratio = Sensible
heat loss/Evaporative
heat loss

Changes in photosynthesis as a function of temperature

A temperature range where enzymatic events are stimulated;

A range of optimal photosynthesis;
A range where destructive events occur.

The relative rates of photosynthetic carbon gain

Southern Texas

Manitoba, Canada

Reduced quantum yield and increased photorespiration

leads to differences in the photosynthetic capacities of C 3
and C4 plants at different latitude

Concentration of atmospheric CO2 from 420,000 years ago to present

Atmospheric CO2 levels have been increasing since the

industrial revolution due to human use of fossil fuels.

Points of resistance to the diffusion of CO2

Concentration
gradients drive the
diffusion of CO2 from
the atmosphere to
rubisco, using both
gaseous and liquid
routes.
The stomatal pore is
the major point of
resistance to CO2
diffusion

Distribution of absorbed light; Spinach leaf cross section (Part 1)

Distribution of absorbed light in spinach sun leaves (Part 2)

Within the leaf depleted of blue and red photons,

green light penetrates deeper into the leaf and
effectively supplies energy for photosynthesis

Atmospheric carbon dioxide levels and daytime growing season temperatures

C4 photosynthesis may have become prominent in warmest regions

when global CO2 concentrations fell below a threshold value;
C4 plants can use water and nitrogen more efficiently than C3 plants
can; but on the other hand, the cost for concentrating CO 2 makes it
less efficient in light utilization.

Photosynthetic carbon assimilation, evaporation, and stomatal conductance

CAM plants

CAM plants can survive severe drought stress. CAM idling: stomata
are closed all the time, CO2 released by respiration is refixed into
malate under severe drought stress.

Stable carbon isotope in plants

C12 and C13 are stable carbon isotopes in the air

R = 13CO2 /12CO2
The carbon isotop ratio of plants

13C (%o) = (Rsample/Rstandard -1) *1000

Rstandard: carbonate of belemnite as standard
13C:
C3 plants: -28 %o
C4 plants: -14%o
Air:
-8%o

Frequency histograms for observed carbon isotope ratios in C3 and C4 taxa

Carbon isotope

Why are there carbon isotope ratio variation in

plants?
12CO2 is lighter and diffuses faster than 13CO2;
Rubisco has an intrinsic discrimination against
13C (~ -30%o); PEP carboxylase has a much
smaller isotope discrimination effect (~ 2%o).
When stomata is more open in C3 plants, more
discrimination of C13. C3 plants grown under
water stress conditions tend to have more
positive (less negative) carbon isotope ratio.

Vegetation changes occur along rainfall gradients in Australia

Increased rainfall results in increased carbon

discrimination (higher negative value)

CAM carbon isotope

CAM plants have similar 13C values to those of C4

plants
For facultative CAM plants, when well watered, it
could use C3 pathway to fix CO2. Thus, the 13C
values of CAM plants reflect how much carbon is
fixed via the C3 pathway versus the C4 pathway