Principles of terrestrial ecosystem ecology.pdf

from diurnal variations in photosynthetic
carbon gain.
Organisms adjust seasonally in response to
changing photoperiod (day length). Many temperate
plants, for example, exhibit a relatively
predictable pattern of phenology, the seasonal
timing of production and loss of leaves, flowers,
fruits, etc. Plant leaves begin to senesce and
reduce their rates of photosynthesis when day
length or other environmental cues signal the
characteristic onset of winter. Before senescence,
plants transport carbohydrates and
nutrients from leaves to storage organs to
prevent their loss during senescence (Chapin et
al. 1990). These stores provide resources to
support plant growth the next spring, so NPP
does not depend entirely on acquisition of new
resources at times when no leaves are present.
Other ecosystem processes change as either
direct consequences of changes in environment
(e.g., the decline in decomposition during
winter due to lower temperatures) or indirect
consequences of changes in other processes
(e.g., the pulse of litter input to soil after leaf
senescence). Ecosystem processes largely
recover after each period of the cycle due to the
predictable nature of diurnal and seasonal perturbations
and the resilience of most processes
to these changes. It is therefore unnecessary to
consider explicitly the physiological basis of circadian
and photoperiodic controls to predict
ecosystem processes over longer time scales
(see Chapter 13). In contrast to temperate
ecosystems, tropical wet forests exhibit a less
well-defined seasonality. Individual species frequently
shed their leaves synchronously, but
species differ in their timing of senescence,
so the ecosystem as a whole shows no strong
seasonal pulse of production and senescence.
The seasonality of plant growth depends on
the seasonality of leaf area and factors regulating
photosynthesis. Spring growth of plants is
initially supported by stored reserves of carbon
and nutrients that were acquired in previous
years. Leaves quickly become a net source of
carbon for the rest of the plant, and growth
during the remainder of the growing season is
largely supported by the current year’s photosynthate.
There is often competition among
Allocation 135
plant parts for allocation of a limited carbohydrate
supply early in the growing season,
resulting in a seasonal progression of production
of different plant parts, for example, with
leaves produced first, followed by roots, and
then by wood (Kozlowski et al. 1991). Plants
species differ, however, in their seasonal patterns
of allocation. Plants with evergreen leaves
may allocate NPP to root growth earlier than
would deciduous plants, because they already
have a leaf canopy that can provide carbon
(Kummerow et al. 1983). Ring-porous temperate
trees must first allocate carbon to xylem
production in spring to develop a functional
water transport system. The water columns
in their large-diameter vessels cavitate (break)
during winter freezing, so xylem vessels
remain functional for only a single growing
season. This large carbon requirement to
rebuild xylem vessels each spring may explain
the northern boundary of ring-porous species
such as oaks (Zimmermann 1983). Seedlings
in dry environments often depend entirely on
their cotyledons for photosynthesis during the
first weeks of growth and allocate all NPP to
root growth to provide a dependable water
supply. The allocation calendar of a plant provides
a general seasonal framework for allocation.
Variations in environment cause plants
to modify this allocation calendar to achieve
the appropriate balance of carbon, water, and
nutrients.
In ecosystems with short growing seasons,
such as arctic tundra, a substantial proportion
of the “current” year’s production is actually
supported by resources that were acquired in
previous years. In late summer, carbon and
nutrient stores are replenished to support the
next year’s production. This seasonal pattern of
storage buffers plant production from seasonal
and interannual variations in the environment
(Chapin et al. 1990). The seasonality of plant
growth is constrained by the availability of leaf
area early and late in the growing season but
otherwise follows seasonal patterns of factors
that govern photosynthesis (temperature, light,
and moisture); the relative importance and
timing of these seasonal controls differ among
species and ecosystems.