2
Photosynthesis, Respiration, and
Long-Distance Transport
2A. Photosynthesis
1. Introduction
Approximately 40% of a plant's dry mass consists
of carbon, fixed in photosynthesis. This process is
vital for growth and survival of virtually all plants
during the major part of their growth cycle. In fact,
life on earth in general, not just that of plants, totally
depends on current and/or past photosynthetic
activity . Leaves are beautifully specialized organs
that enable plants to intercept the light necessary
for photosynthesis. The light is captured by a large
array of chloroplasts that are in close proximity to
air and not too far away from vascular tissue , which
supplies water and exports the products of photo-
synthesis. CO2 uptake occurs through leaf pores,
the stomata, which are able to rapidly change their
aperture (see Sect. 5.4 in plant water relations) .
Once inside the leaf, CO2 diffuses from the intercel-
lular air spaces to the sites of carboxylation in the
chloroplast (C3 species) or in the cytosol (C4 and
CAM species).
Ideal conditions for photosynthesis include an
ample supply of water and nutrients to the plant,
as well as optimal temperature and light condi-
tions . Even when the environmental conditions
are less favorable, however, such as in a desert
or the understory of a forest, photosynthesis-
at least of the adapted plants-eontinues. This
chapter addresses how such plants manage to
continue to photosynthesize and/or protect
their photosynthetic machinery in such adverse
environments.
10
H. Lambers et al., Plant Physiological Ecology
Springer Science+Business Media New York 1998
2. General Characteristics of the
Photosynthetic Apparatus
2.1 The Light and Dark Reactions
of Photosynthesis
The primary processes of photosynthesis occur
in the chloroplast. In C3 plants most of the
chloroplasts are located in the mesophyll cells of
the leaves (Fig. 1) . Three main processes are
distinguished:
1. Absorption of photons by pigments, mainly
chlorophylls, associated with two photosystems.
The pigments are embedded in internal membrane
structures (thylakoids) and absorb a major part of
the energy of the photosynthetically active radia-
tion (PAR; 400 to 700nm) . They transfer the exci-
tation energy to the reaction centers of the
photosystems where the second process starts.
2. Electrons derived from the splitting of water
with the simultaneous production of oxygen are
transported along an electron-transport chain em-
bedded in the thylakoid membrane. NADPH and
ATP produced in this process are used in the third
process. Because these two reactions depend on
light energy, they are called the light reactions of
photosynthesis.
3. The NADPH and ATP are used in the photo-
synthetic carbon-reduction cycle (Calvin cycle), in
which CO 2 is assimilated, leading to the synthesis
of C3 compounds (triose-phosphates). This process
General Characteristics of the Photosynthetic Apparatus 11

~--s

FIGURE 1. Above: Scanning electron micros-

cope cross-sectional view of a leaf of Nicotiana
tabacum (tobacco) that shows palisade tissue be-
neath the upper (adaxial) epidermis and spongy
tissue adjacent to the (lower) abaxial epidermis.
Below: Transmission electron microscope micro-
graph of a tobacco chloroplast that shows stacked
(grana, g) and unstacked regions of the thylakoids,
stroma (s), and starch granules (sg). Note the close
proximity of two mitochondria (m, top and bot-
tom) and one peroxisome (p) (scale bar is 111m)
(courtesy I.R. Evans, Research School of Biolog-
ical Sciences, Australian National University,
Canberra, Australia).