Ecosystem Ecology general points

(1) Biotic community is truly inseparable from the abiotic

environment
(2) Energy flow through communities ultimately stems from
the sunlight that is assimilated by plants (Net Primary
Productivity) and becomes available to the remainder of the
food web
(3) Each step along the way, energy is lost through inefficient
transformation mediated by biotic factors
(4) That is the biological details (i.e., species identity or
functional group) greatly impacts E-flow through ecosystems

Cont
(5) Nutrient availability mediates E-flow through ecosystems
because nutrients NOT energy may be most limiting
(6) Nutrient (water, C, N, P) flow through ecosystems is
largely under control of large-scale geological processes
(7) But, the role of individual organisms/species, while often
trivial in terms of global impact, is extremely important in terms
of its impact on the biological world

The Ecosystem Concept A brief history

Early 1910-1920s:
Biotic descriptions
of communities

1935
Include abiotic factors

Clements and Gleason

views plant communities

as a superorganism

emphasized individualistic
properties a fortuitous collection
occurring together in time and space

The Ecosystem Concept A brief history

1927
Emphasized
energy flow

Charles Elton

characterized feeding relationships among

animals on Bear Island, North Atlantic

coined the terms food chain and

food cycle to be replaced by food web

Eltons Pyramids
As one goes up the food chain, one ascends a progression of
sizes as most predators consume smaller prey; larger animals
require more space to find food and hence their numbers go down.

#s
1
10
100
1000

Biomass
1kg
10
100
1000

Of the total solar radiation, 0 , plants

use some fraction, 1 , herbivores
assimilate less energy, 2 , owing to
plants maintaining themselves before
being eaten and E lost during assimilation

2
1

1942

= Biological Efficiency
of the trophic link (<<1)

Ray Lindeman

visualized a Pyramid of Energy Transformation

less energy becomes available to higher levels
due to work being done and inefficient transfer.

1953 Eugene Odum model Energy flow, later adapted for nutrients as well
(1) Energy flows in one direction, absorbed light is lost as heat or transferred
into chemical energy through photosynthesis (Annual Gross Primary
Production) by autotrophic organisms.
(2) Autotrophs spend some energy to respire, other goes into growth, which
becomes available to heterotrophs at the next higher level (Annual Net
Primary Production)

Later, nutrient flows were added:

Unlike energy, nutrients are typically
retained and regenerated in biological
systems. There are some inputs and
outputs (but rather small) and nutrients
pool in some parts of the ecosystem,
but mainly are transferred between
various trophic levels.

 Cycling of nutrients has assumed a near equal status with energy flow.
One reason is that the amounts of the elements and their movement
between ecosystem components provides an index to energy flow, which
may be otherwise difficult to measure.
Carbon in particular bears a close relationship w/energy because of its
intimate association w/photosynthesis.
Also, certain nutrients regulate Primary Production
e.g., water in deserts, N & P in oligotrohic lakes

ENERGY FLOW

Photosynthesis:
6CO2 +6H2O C6H12O6+ 6O2

Or cutting, drying, and weighing

of all plants at the end of the growing
season Annual Aboveground Net
Productivity (AANP).

Photosynthesis How efficient?

Light, temp, water, and nutrients all limit photosynthesis.
Plants are rarely at max (cloud cover, shading)
Rates saturate at high temp (i.e., diminishing returns)
Water loss via stomata can shut down PhotoS (Transpiration efficiency ~ 2g/kg water).
For the ecosystem, efficiency is
~ 1-2%. Plants reflect 25-75% of
incoming light and another large
percent is absorbed by other molecules
and lost as heat.
Finally, nutrient availability affects
plant productivity

C3

C4 CAM

Because ecosystems differ in terms

of plant density, water input and
availability, nutrient availability,
and the uniqueness of individual spp
adapted to particular environments,
Annual Primary Productivity varies
greatly with ecosystems.

Energy transfer between trophic levels is only 5-20% efficient

Ingestion
Egestion
Energy
For Excretion
Detritus (N waste)
feeders

Digestion &
Assimilation
Growth and
Reproduction

ingested E egested E
= assimilated E
(mainly a function of diet)

Respiration, energy
used to perform work
is lost
assimilated E respiration
= production
(mainly a function of metabolic rate)

Death

Next level
Ecological Efficiency is the ratio
of biomass at one trophic level that is
incorporated into biomass at the next trophic level.

1-6% active, endotherms

up to 75% for aquatic species
plants: 30-85%

Detritus then, often dominates

food webs at least in terrestrial
systems.

Herbivory rates
Temp forest: 2-3%
Old fields: 12%
Lakes: 60-99%

Food chain lengths are ultimately limited by ecological efficiencies

Community

NPP

Ingestion

Eff

(kcal/m /yr)

(kcal/m2/yr)

(%)

500
8000
2000
8000

0.1
10
1.0
10

25
20
10
5

Open ocean
Coastal marine
Temp. grassland
Tropical forest

n
7.1
5.1
4.3
3.2

(5.5)
(6.6)
(4.3)
(4)

Pathways of Elements in the Ecosystem a generalization

Ecosystem is broken down

into separate compartments (in
a hierarchical manner) or pools
where elements reside.
Movement between pools often
biochemical transformation,
often involving energy.
Photosynthesis adds energy to
carbon, whereas burning fossil
fuels removes energy.

**Responsible for the

imbalance in the
modern C-cycle

New**

Global Carbon Cycle

3 classes of cycling through
aquatic and terrestrial systems:
(1) Assimilation/dissimilation
reactions via photosynthesis
and respiration
(2) C02 exchange between the
oceans and atmosphere
(3) Sedimentation of carbonates

#s refers to Billions of
metric tons - 1015 grams

Global Nitrogen Cycle

** in 2000 anthropogenic sources of N exceeded natural sources
at 87 x 106 MT forecast for 2020: 135 MMT; 2050: 236 MMT
(pesticides - 2000: 3.75 MMT, 2050: 10.1 MMT)

**

Precip
+ 76 kg/ha/yr

E.g., of Local Nitrogen Cycle

Model of N-cycling in a forest
Undisturbed forest
gains 1-3 kg/ha/yr

1200 Kg/ha

Soil
14,000 Kg/ha

Tropical Forest
in Ghana: 1794/4587

Model of N-cycling in a forest

(disturbed Forest)
Precip
+ 76 kg/ha/yr

Fire (wind)

Clear-cut
- 54kg/ha/yr
Large blow-downs
Defoliating insects???
- recovery is quick
- trees remain alive
- no soil erosion
Soil

Insect defoliation and N-cycling in forests

Gypsy moths are an introduced
insect pest that experiences
occasional outbreaks in which
huge swaths of oak forest are
entirely defoliated

Fate of Foliar N (15N-labeled exp)

Normal
Resorption (fall)
Fall leaf drop
Early green leaf drop
Leaching
Insect Biomass
Frass

70%
25
1.4
2
0.7
1

Outbreak
23
17 (new)
29
4
8
23

-47
-8
+28
+2
+7
+22

N has very different fates Frass/insects biomass

at the expense of resorption

What is the fate of foliar N consumed by insects and

deposited as green fall??
<0.1% Gaseous N
Assimilated: 16%

Frass: 84% N
100% N

Frass: Scatological Research Team 9% organically extractable

91% salts, such as uric acid
Frass is readily consumed/broken down by microbes,
which immobilizes N in microbial biomass
Oak litter: 81% N recovered, mostly as undecomposed
leaf litter.
Frass: 40% recovered, almost all of it through the soil

But, defoliation disrupts normal N-cycling through

soil-plant systems
Litter N is released from leaf litter slowly via decomposition,
whereas
Frass N moves quickly into the subsoil where it is retained as
organic matter in mircobial biomass and released more slowly
into the soil pool of N.
As a consequence, short-term effects of Gypsy moth outbreaks
have a small effect on N-cycling through the forest, however,
chronic outbreaks/defoliation can lead to heavy leeching,
depletion of soil N and high concentration of NO3 and NO2

Punch line species are intimately tied to the cycling and transfer
of Energy and Nutrients through ecosystems.
E.g., #1 Photosynthetic rates
E.g., #2 Assimilation rates/ecological efficiencies
E.g., #3 Impacts of gypsy moths on N-cycling
Will the loss of species richness from communities severely
impact ecosystem processes ?

Will the loss of species richness form communities severely

impact ecosystem processes ?
3 Hypotheses

redundant

Ecosystem
Process/service
e.g., productivity,
pollination

idiosyncratic

Non-substitutable
Species richness

Ecosystem
Process, e.g.,
productivity

Species richness
Non-substitutable implies that every species is unique and its
Effects/impacts cannot be substituted by others.

Ecosystem
Process, e.g.,
productivity

Species richness
Redundancy implies that every species can partially compensate
for each others absence, and that there is diminishing returns to
species diversity in terms of their ability to supply ecosystem
services i.e., not all species are necessary for a fully functional
ecosystem

Ecosystem
Process, e.g.,
productivity

Species richness
Idiosyncratic implies that each unique combination of species
has its own level of ecosystem service provision and is not
predictable a priori

Why should there be a positive correlation be ecosystem

service and species richness?
Ans #1: Sampling artifact. By chance, a larger group of species
is more likely to contain the MOST productive species than a
small group.

Never exhibits over-yielding, that is, no group of species

can ever outperform a monoculture of the best single species
Productivity

group

A, B, C individually

Why should there be a positive correlation be ecosystem

service and species richness?
Ans #2: Niche- complementarity competition theory:
species will evolve to be distinct and thereby reduce
or eliminate competition.

e.g., root
profiles

Because a group of species can more effectively harvest water,

productivity of the group is enhanced over any single species
C
B
Vs.
A

Productivity

Exhibits over-yielding

group

A, B, C individually

Decomposition of leaf packs in Williamstown, MA

.3
.2
K-value

(decomposition rate) .1

0
-.1
Lonicera maple

beech hemlock

The implication is that a world dominated by Lonicera

releases large amounts of organic matter quickly, but
little remains for the end of the growing season. Therefore,
diversity/richness of stream invertebrates is reduced in
systems with reduced canopy cover
nutrient availability

time
Lonicera maple

beech hemlock

Species diversity/richness and temporal variability

Two lines of thought:
(1) Insurance Hypothesis rare species are extremely
important under extreme conditions.
(2) Portfolio Effect More diverse portfolios are less volatile
because, several randomly and independently variable
items are less variable than the average item
Services provide by the groups of species include:
stability (the inverse of variability), function during
extreme events (e.g., drought resistance), ability to
resist invasion

Cedar Creek LTER (Univ. MN David Tilman et al.)

Plant biomass (% cover)

and soil denitrification
rates (less soil nitrate)
increase with species
richness

Plant biomass shows greater

resistance to drought when
species richness is higher
Rare spp response

Loss of species is a concerned because the Earths biota

performs many valuable functions and services

Valuing species richness and diversity

Aesthetic
Research and Development
Economical
Service (and $$$)