Atmosphere/biosphere interactions: Primary productivity as an example

Outline:
Introduction
Review of photosynthesis
o Water-use efficiency in photosynthesis
! Definition of stable isotopes
o Nutrient-use efficiency
Respiration
Net Primary Production
Fate of Net Primary Production
o Production of detritus
o The decomposition process
o Humus formation and soil organic matter

Please note that the majority of the text below is taken from Chapter 5 in:
Schlesinger, W.H. 1997. Biogeochemistry: an analysis of global change. 2nd edition. Academic Press, California.

Photosynthesis is biogeochemical process that transfers carbon from oxidized form (CO2) in
atmosphere to reduced form (organic) in the biosphere.
Photosynthesis provides the energy for all other forms of life in biosphere
Plant growth affects composition of:
o atmosphere (e.g., O2, CO2)
o soil development

Net Primary Productivity: rate of accumulation of organic matter in tissues of plants

Subsequent storage of organic matter in:
o Plant tissues (biomass)
o Dead plant parts (detritus)
o Soil organic matter

In addition to CO2, light and water, plants require nutrients for photosynthesis
Nutrients ultimately derived from atmosphere or underlying bedrock

Overall storage of carbon on land results from balance between net primary production and
decomposition (which returns CO2 back to atmosphere)

Review of Photosynthesis

Chlorophyll contains central atom of Mg (example of how abundant product of rock weathering is
essential element in biochemistry)

When photosynthetic pigments absorb sunlight, a few chlorophyll molecules are oxidized
Passes electron to sequence of electron transfer proteins which ultimately reduce a high
energy molecule NADP to NADPH

Chlorophyll molecule regains electron from water, which is split by enzyme containing Mg, Ca and
Cl
Basically: 2H20 4H+ + 4e- + O2
 o Protons then used to synthesize another high-energy protein: ATP

NADPH and ATP then used by suite of enzymes (e.g., Rubisco) to reduce CO2 and build
carbohydrates
Overall reaction: CO2 + H20 CH2O + O2

NOTE: O2 released to atmosphere comes from H20 not CO2

Rate of photosynthesis controlled by stomatal conductance

Stomatal conductance controlled by availability of H2O and concentration of CO2 inside leaf (which
is consumed by photosynthesis)

Water use efficiency

When stoma are open, H20 can diffuse outward
In Hubbarb Brook Experimental Forest (New Hampshire), ~25% of annual precipitation
lost by plant uptake and transpiration

WUE = mmoles CO2 fixed/moles H2O lost

Water use efficiency higher at lower stomatal conductance
Rising CO2 concentrations in atmosphere due to human activities may have resulted in:
1. Same rate of photosynthesis at lower stomatal conductance
2. Lower number stomates per unit of leaf surface

LONG-TERM water use efficiency may be estimated from carbon isotope composition of plant
tissues

Definition of stable isotope

A chemical element's atomic number is the number of positive charges (protons) in the nucleus
This number is invariant for all atoms of that element

If some atoms of an element have a different atomic weight from others, the difference is in the
number of neutrons
Atoms of the same atomic number but different atomic weights are called isotopes.

Elements can exist in both stable and unstable (radioactive) forms

Most elements of biological interest (including C, H, O, N, and S) have two or more stable
isotopes, with the lightest of these present in much greater abundance than the others

Among stable isotopes the most useful as biological tracers are the heavy isotopes of carbon and
nitrogen
These two elements are found in the earth, the atmosphere, and all living things
o Each has a heavy isotope (13C and 15N) with a natural abundance of ~1% or less and
a light isotope (l2C and 14N) that makes up virtually all the remainder

Average Terrestrial Abundances of the Stable Isotopes of

Major Elements of Interest in Ecological Studies
Element Isotope Abundance (%)
Hydrogen 1H 99.985
2H 0.015
Carbon 12C 98.89
13C 1.11
Nitrogen 14N 99.63
15N 0.37
Oxygen 16O 99.759
17O 0.037
18O 0.204
Sulfur 32S 95.00
33S 0.76
34S 4.22
36S 0.014

Method based on observation that diffusion of 12CO2, a lighter molecule, is more rapid than that of
13CO2 (which composes ~1.1% of the atmosphere)

Therefore more 12CO2 enters the leaf than 13CO2

o Once in leaf, Rubisco also has higher affinity for 12CO2
! As a result, plant tissue contains 2% less 13CO2 than the atmosphere (or 20
)

This fractionation is mathematically expressed relative to an accepted standard as follows:

13C = ((13C/12Csample 13C/12Cstandard)/(13C/12Cstandard))*1000

Using the units of parts per thousand ()

Because atmospheric CO2 shows an isotopic ratio of 8 versus the standard, most plant tissues
show 13C of ~ -28 [i.e., (-8) + (-20)].

Discrimination between 12CO2 and 13CO2 during photosynthesis is greatest when stomatal
conductance is high (Figure 2.6)
When stomates are partially or completely closed, nearly all CO2 in leaf reacts with Rubisco,
and there is less fractionation of the isotopes
o Therefore isotopic ratio of plant tissue directly related to average stomatal
conductance during growth

13C values of preserved plant materials indicate that water use efficiency of plats has increased as
concentration of atmospheric CO2 rose:
1. at the end of the last glacial period
2. during past several hundred years

Nutrient Use Efficiency

Rate of photosynthesis is directly correlated to leaf nitrogen content on a mass basis (Figure 5.3)
Rubisco accounts for ~20-30% of leaf nitrogen, and therefore photosynthesis may be
limited by enzyme production
o Phosphorus may also be important determinant of photosynthesis
o Despite central role, Mg and Mn are seldom limited

Variations in nutrient use efficiency are seen among:

different plant types
 plants grown at different fertility levels

For many plant species, when leaf nutrient content increase (e.g., via fertilization), nutrient use
efficiency declines
Nutrient use efficiency also appears inversely correlated to water use efficiency across many
species

Respiration

Net photosynthesis: fixation of carbon in excess of the simultaneous release of CO2 by plant metabolism
(respiration)
Correlated with nitrogen content of plant cells (Figure 5.4)

In woody plants, large fraction of respiration contributed by stems and roots owing to their large
contribution to plant biomass
For leaf tissues, respiration higher in day than at night due to process of photorespiration

~50% of gross carbon fixation is used by plants

for long-lived woody plants, maintenance respiration increases with age resulting in slower
growth
respiration generally increases with temperature (high rates of respiration n tropics, and
possibly due to global warming)

Net Primary Production

gross primary production (GPP) plant respiration (Rp) = net primary production (NPP)

NPP not equivalent to plant growth

Some loss to:
herbivores
death and loss of tissue (litterfall)

HOW DO WE MEASURE NPP?

Allocation of NPP varies with:

Vegetation type
Age

In forests
25-35% of aboveground production found in leaves (decreases with stand age)
In shrublands
35-60% allocated to foliage
In grasslands
almost 100% NP found in photosynthetic tissues

When comparing plant communities:

% allocation of NPP to wood growth greater in boreal forests than tropical forests
o because of massive structure and high temperatures, tropical forests may expend
more GPP in respiration
Figure 5.4

Root growth difficult to quantify

However, when it has been examined, annual growth and turnover of root tissues account
for significant fraction of NPP in most communities
In forests:
o proportion NPP allocated to root growth inversely related to soil fertility
o absolute amount of root growth greatest on sites with high NPP

Table 5.1

In forests:
Typically during the day, there is a net uptake of CO2 from the atmosphere
During the night, the gradient is often reversed as plant and soil respiration continue in
absence of vegetation uptake

Often measured with the eddy correlation technique

e.g.,
In Massachusetts deciduous forest:
o GPP: 1070-1210 g C m-2 yr-1
o Plant and soil respiration: 810-1140 g C m-2 yr-1
o Net accumulation of carbon in ecosystem: 140-280 g C m-2 yr-1

The Fate of NPP

Plant communities achieve steady state in living biomass when allocation to woody tissue is balanced
by death and loss of older parts (Figure 5.13)
At that point, there is no true increment in biomass, although dad organic matter may still be
accumulating in soil.

Net Ecosystem Production (NEP)

NEP = NPP (Rh + Rd)

Where:
NPP = GPP - Rp
Rh = herbivore respiration
Rd = decomposer respiration

Therefore:

NEP = GPP Rt

Where Rt is the total respiratory loss of CO2 from the ecosystem.

Figure 5.14

Relationships suggest that increments in organic matter possible only during early stages of plant
community development
In older communities, there is no true increment to live biomass, and nearly all the NPP is
delivered to soils, where it decomposes
 o NOTE: role of animals is relatively minor
o The consumption of plant tissues by herbivores is nearly always <20% of NPP
! By consuming leaves and roots, however, herbivores may have indirect
effect on NPP than estimate above.

Fires analogous to respiration by herbivores and decomposers

Rapidly converts long-term accumulation of NEP to CO2 and restarting ecosystem
development

Production of Detritus:

Largest fraction of NPP is delivered to soil as dead organic matter

Global patterns in decomposition of plant litterfall similar to global patterns in NPP
o Deposition of litterfall declines with increasing latitude from tropical to boreal
forests

Leaf tissues account for ~70% of aboveground litterfall in forests

Decomposition of woody litter tends to increase with forest age
o Fallen logs conspicuous component of forest floor in old-growth areas

In grassland ecosystems, annual litterfall nearly equal to annual NPP

In most areas, annual growth and death of fine roots contributes large amount of detritus to soil
(often overlooked)

The Decomposition Process

Most detritus is delivered to upper layers of soil where it is subject to decomposition by microfauna,
bacteria and fungi
Decomposition leads to release of CO2, H2O and nutrient elements
Also have microbial production of highly resistant organic compounds known as humus
o Humus accumulates in lower soil profile and compose bulk of soil organic matter

Pool of carbon in soils result of two stages of decomposition:

Processes leading to rapid turnover of majority of litter a surface
Processes leading to slower production, accumulation and turnover of humus at depth

Using a mass-balance approach, annual decomposition should equal annual input of fresh debris, so
mass of detritus stays constant.
Under these assumptions, a constant fraction (k) of the detrital mass decomposes:

Litterfall = k(detrital mass) or (litterfall)/(detrital mass) = k

When decomposition rates are rapid, there is little surface layer accumulation (k > 1.0)
e.g., in the tropics, where decomposition has potential to respire more than annual input of
carbon in litterfall

In contrast, in peatlands values for k are very small (e.g., 0.001)

Decomposition in grasslands shows range of k of 0.2 to 0.6

Values for deserts may be as high as 1 due to:
Action of termites
Photooxidation of litter by UV

Global mean residence time of 3 years (k=0.33) for carbon on surface of soil

Decomposition rates vary as function of:

Temperature (microbial activity increases exponentially with increasing temperature)
Moisture (limits decomposition in arid and semiarid regions) (Figure 5.15)
Chemical composition of litter material

Humus Formation and Soil Organic Matter

Plant litter and soil microbes constitute cellular fraction of soil organic matter
As decomposition proceeds, there is increasing content of non-cellular organic matter or
humus, resulting from microbial activity

Under most vegetation, mass of humus in soil profile exceeds combined content of organic matter in
forest floor and aboveground vegetation (Table 5.3)

Global estimate of soil organic matter DIVIDED BY estimate of global litterfall suggests mean
residence time of ~30 years for total pool of organic carbon in soils
Varies over several orders of magnitude between surface litter and various humus fractions
(Figure 5.17)

Global distribution of soil organic matter shows how moisture and temperature control balance
between primary production and decomposition in surface and lower soil layers (Table 5.3)
Accumulation of soil organic matter:
o Greatest in wetland ecosystems
o Least in deserts

Accumulations increase from tropical to boreal climates

NPP shows opposite trend, so accumulation largely due to differences in decomposition

Worldwide, accumulation of soil organic matter seems more related to factors controlling
decomposition than to NPP of terrestrial ecosystems.

Storage of soil organic matter represents the net ecosystem production (NEP) in terrestrial
ecosystems.
Although many wetland ecosystems show long-term net accumulations, mass of soil organic
matter in most upland ecosystems has been fairly constant before widespread human
disturbance of soils (Table 5.4)

NOTE:
Total storage of carbon in soils can only account for 0.03% of O2 content of atmosphere, given that
the storage of organic carbon and the release of O2 occur on a mole-for-mole basis during
photosynthesis
Thus accumulation of atmospheric O2 cannot be result of storage of organic C on land
o Long-term storage of organic C appears to be dominated by accumulation in anoxic
marine sediments