Chapter 8: Nutritional Regeneration in

Terrestrial and Aquatic Ecosystems

Robert E. Ricklefs
The Economy of Nature, Fifth Edition

(c) 2001 by W. H. Freeman and

Company
Acid Rain and Forest
Growth

Decline in forests, noted in northeastern

US and central Europe in the 1960s,
appeared correlated with acid rain.
The Clean Air Act of 1970 reduced
emissions of sulfur oxides and particulates
in the US.
Forests did not show signs of recovery.
Why?

(c) 2001 by W. H. Freeman and

Company
Slow Recovery of Forests
from Effects of Acid Rain

Studies at Hubbard Brook Experimental

Forest in New Hampshire showed why
forests did not recover after passage of
Clean Air Act:
acidity of rain declined slowly
emissions of particulates declined, reducing an
important source of calcium at Hubbard Brook
leaching of calcium and other nutrients by acid
rain left lasting effects on soil fertility
(c) 2001 by W. H. Freeman and
Company
Lessons from Hubbard
Brook

Acidity itself is not the cause of tree

death:
long-term leaching of nutrients kills trees
Natural recovery will be slow on nutrient-
poor soils:
restoration of nutrients will require
weathering
weathering is a slow process
(c) 2001 by W. H. Freeman and
Company
More Lessons from
Hubbard Brook

Effects of acid rain on soils may remain

for years, even if causes of the problem
are addressed.
Understanding nutrient cycling and
regeneration is crucial to understanding
ecosystem function.

(c) 2001 by W. H. Freeman and

Company
Nutrient regeneration
occurs in soils.

Nutrients are added to the soil through

weathering of bedrock or other parent
material.
How fast does such weathering occur?
estimates can be made for positive ions such
as Ca2+, K+, Na+, and Mg2+
at equilibrium, net losses must be balanced
by replenishment from weathering
(c) 2001 by W. H. Freeman and
Company
Weathering of Ca2+ at
Hubbard Brook

Watershed budgets:
precipitation inputs = 2 kg ha-1 yr-1
streamflow losses = 14 kg ha-1 yr-1
assimilation by vegetation = 9 kg ha-1 yr-1
net removal thus = 21 kg ha-1 yr-1
Total weathering of bedrock to offset Ca+2
losses is 1,500 kg ha-1 yr-1 or 1 mm depth.
Later analyses showed this to be an
overestimate; the system was not in equilibrium.

(c) 2001 by W. H. Freeman and

Company
Quality of detritus influences the
rate of nutrient regeneration.

Weathering is insufficient to supply plants

with essential elements (Ca, Mg, K, Na, N,
P, S, etc.) at the rates required.
Rapid regeneration of these elements
from detritus is essential for ecosystem
function.
In forests, detritus is abundant:
includes plant debris, animal excreta, etc.
>90% of plant biomass
(c) 2001 enters
by W. H. Freeman and detritus pool
Company
Breakdown of Leaf Litter

Breakdown is a complex process:

leaching of soluble minerals:
10-30% of substances in leaves are water-soluble
consumption by large detritivores:
assimilate 3-40% of energy
macerate detritus, speeding microbial activity
breakdown of woody components by fungi
decomposition of residue by bacteria
(c) 2001 by W. H. Freeman and
Company
Quality of Plant Detritus

Litter of various species decays at different rates:

weight loss in 1 yr for broadleaved species varied from
21% for beech to 64% for mulberry
needles of pines and other conifers decompose slowly
resistance to decay is largely a function of
composition, especially lignins, which resist decay
Fungi play special roles in degrading resistant
materials:
fungi especially capable of degrading cellulose, lignins

(c) 2001 by W. H. Freeman and

Company
Mycorrhizae

Mycorrhizae are mutualistic associations

of fungi and plant roots:
endomycorrhizae - fungus penetrates into
root tissue
ectomycorrhizae - fungus forms sheath
around root
Mycorrhizae facilitate nutrient extraction
from nutrient-poor soils, enhancing plant
production.
(c) 2001 by W. H. Freeman and
Company
Function of Mycorrhizae

Mycorrhizae are effective at extracting nutrients:

penetrate larger volume of soil than roots alone
secrete enzymes and acids, which extract nutrients
Endomycorrhizae are associated with most
plants:
apparently an ancient association
fungi are specialists at extracting phosphorus
Ectomycorrhizae are also widespread:
sheath stores nutrients and carbon compounds
fungi consume substantial amount of net production
(c) 2001 by W. H. Freeman and
Company
Climate and Nutrient
Regeneration

Nutrient cycling is affected by climate:

temperate and tropical ecosystems differ
because of effects of climate on:
weathering
soil properties
decomposition of detritus
In temperate soils, organic matter provides
a persistent supply of mineral elements
released slowly by decomposition.
(c) 2001 by W. H. Freeman and
Company
A Tropical Paradox

Tropical forests are highly productive in spite of

infertile soils:
tropical soils are typically:
deeply weathered
have little clay
do not retain nutrients well
high productivity is supported by:
rapid regeneration of nutrients form detritus
rapid uptake of nutrients
efficient retention of nutrients by plants/mycorrhizae

(c) 2001 by W. H. Freeman and

Company
Slash-and-Burn Agriculture

Cutting and burning of vegetation initiates

the cycle:
nutrients are released from felled and burned
vegetation
2-3 years of crop growth possible
fertility rapidly declines as nutrients are
leached
upward movement of water draws iron and
aluminum oxides upward, resulting in laterite
(c) 2001 by W. H. Freeman and
Company
Is Slash-and-burn
sustainable?

Traditional agriculture is sustainable:

2-3 years of cropping depletes soil
50-100 years of forest regeneration rebuilds
soil quality
Population pressures lead to acceleration
of the cycle:
soils are insufficiently replenished
soils deteriorate rapidly, requiring expensive
fertilizer subsidies
(c) 2001 by W. H. Freeman and
Company
Vegetation and Soil
Fertility

Vegetation is critical to development and

maintenance of soil fertility:
clear-cutting of an experimental watershed
at Hubbard Brook, NH resulted in:
several-fold increase in stream flow
3- to 20-fold increase in cation losses
shift from nitrogen storage to massive nitrogen
loss:
• uncut system gained 1-3 kg N ha-1 yr-1
• clear-cut system lost 54 kg N ha-1 yr-1
(c) 2001 by W. H. Freeman and
Company
Soil versus Vegetation
Stocks of Nutrients

Litter and other detritus do not form a

large reserve of nutrients in the tropics:
forest floor litter as percentage of vegetation
plus detritus:
20% in temperate needle-leaved forests
5% in temperate hardwood forests
1-2% in tropical forests
soil to biomass ratio for phosphorus in
forests is 23.1 in Belgium, 0.1 in Ghana

(c) 2001 by W. H. Freeman and

Company
Eutrophic and Oligotrophic
Soils

Tropics have both rich and poor soils:

eutrophic (rich) soils develop in geologically
active areas with young soils where:
erosion is high
rapid weathering of bedrock adds nutrients
oligotrophic (poor) soils develop in old,
geologically stable areas with old soils where:
intense weathering of soils removes clay and
reduces storage capacity for nutrients

(c) 2001 by W. H. Freeman and

Company
Nutrient Retention by
Vegetation

Retention of nutrients by vegetation is

crucial to sustained productivity in tropics.
Plants retain nutrients by:
retaining leaves
withdrawing nutrients before leaves are
dropped
developing dense root mats near soil surface

(c) 2001 by W. H. Freeman and

Company
Nutrients are regenerated
from aquatic sediments.

Soils and aquatic sediments share similar

regenerative processes (both processes
occur in aqueous medium).
Soils and aquatic sediments differ in two
profound ways:
release of nutrients in soils occurs near plant
roots in soils, far from roots in sediments
release of nutrients is aerobic in soils,
anaerobic in aquatic sediments
(c) 2001 by W. H. Freeman and
Company
Nutrients and Aquatic
Productivity

Productivity in aquatic systems is stimulated

when nutrients are in the photic zone, resulting
from
proximity to bottom sediments
upwelling of nutrient-rich water
Regeneration of nutrients by excretion and
decomposition may take place within the water
column.
Sedimentation represents a continual drain on
nutrients within(c)the water
2001 by column.
W. H. Freeman and
Company
Thermal stratification
hinders vertical mixing.

Vertical mixing is critical to replenishment of

surface waters with nutrients from below:
results from turbulent mixing driven by wind
impeded by vertical density stratification:
may be caused by thermal stratification
also occurs when fresh water floats over denser salt water
Vertical mixing has positive and negative effects
on productivity:
nutrients brought from depths stimulate productivity
phytoplankton may be carried below photic zone
(c) 2001 by W. H. Freeman and
Company
Stratification inhibits
production.

Thermal stratification in temperate lakes:

nutrients regenerated in deeper waters cannot reach
the surface
vertical mixing in fall brings nutrient-rich water to the
surface
Stratification in other aquatic systems:
arctic/subarctic and tropical lakes are not thermally
stratified and mix freely
in marine systems, stratified and non-stratified water
bodies may meet, stimulating production

(c) 2001 by W. H. Freeman and

Company
Nutrients limit production
in the oceans.

Primary production of marine ecosystems

is tied closely to nutrient supplies:
nitrogen is especially limiting
shallow seas and areas of upwelling are
especially productive
some areas of open ocean are unproductive,
despite adequate nitrogen and phosphorus:
iron may be limiting in some areas of open ocean
silicon may also be limiting, especially for diatoms
(c) 2001 by W. H. Freeman and
Company
Oxygen depletion facilitates
nutrient regeneration.

Nutrient regeneration is facilitated as anoxic

conditions develop in hypolimnion and
sediments of stratified temperate lakes:
nitrification ceases, leading to accumulation of
ammonia
iron is reduced from Fe3+ to Fe2+
insoluble iron-phosphorus complexes are solubilized,
releasing iron and phosphorus
These processes reverse when oxidizing
conditions return during fall overturn.
(c) 2001 by W. H. Freeman and
Company
Phosphorus and Trophic
Status in Lakes

Phosphorus typically limits productivity in

freshwater systems:
P is especially scarce in well-oxygenated surface
waters
Natural lakes exhibit a wide range of fertilities:
productivity depends on:
external nutrient inputs
internal regeneration of nutrients

(c) 2001 by W. H. Freeman and

Company
Temperate lakes exhibit
varied degrees of mixing.

Productivity depends in part on degree of

mixing of surface and deeper waters:
shallow lakes may lack hypolimnion and circulate
continuously
somewhat deeper lakes stratify sporadically, with
periods of mixing caused by:
strong winds
occasional cold weather in summer
deepest lakes rarely mix completely, so productivity
depends on external nutrient sources
(c) 2001 by W. H. Freeman and
Company
Productivity varies in
temperate lakes.

Lakes may be classified on a continuum from

oligotrophic to eutrophic.
oligotrophic lakes are nutrient-limited and
unproductive
naturally eutrophic lakes exist in a well-nourished
and productive dynamic steady-state
human activities can lead to inappropriate nutrient
loading resulting from:
inputs of sewage
drainage from fertilized agricultural lands
(c) 2001 by W. H. Freeman and
Company
Cultural eutrophication of
lakes is harmful.

Nutrients stimulate primary production.

Production is not inherently harmful, but:
biomass accumulates, overwhelming natural
regenerative processes
untreated sewage also increases the amount
of organic material in water
increased biological oxygen demand
depletes oxygen, killing fish and other
obligate aerobes
(c) 2001 by W. H. Freeman and
Company
Estuaries and marshes are
highly productive.

Shallow estuaries and salt marshes are

among the most productive ecosystems
on earth.
High production in these systems results
from:
rapid and local regeneration of nutrients
external loading of nutrients

(c) 2001 by W. H. Freeman and

Company
Marshes and estuaries
export their production.

Adjacent marine ecosystems benefit from

export of production from marshes and
estuaries. For example,
a Georgia salt marsh exported nearly 50% of
its net primary production to surrounding
marine systems in the form of:
organisms
particulate detritus
dissolved organic material

(c) 2001 by W. H. Freeman and

Company
Marshes and estuaries are critical to
functioning of marine ecosystems.

Marshes and estuaries are important

feeding areas for larval and immature
stages of fish and invertebrates,
providing:
hiding places
high productivity
These organisms later complete their life
cycles in the sea.
(c) 2001 by W. H. Freeman and
Company
Summary

Chemical and biochemical transformations

are modified by physical and chemical
conditions in each type of ecosystem.
Pathways of elements in ecosystems
reflect patterns of nutrient cycling.

(c) 2001 by W. H. Freeman and

Company
Summary: Terrestrial and
Aquatic Systems

In terrestrial systems:

ecosystem metabolism is mostly aerobic
production is limited by regeneration of nutrients from
soils
In aquatic systems:
anaerobic respiration and regeneration of nutrients
occurs in sediments, far from producers
local regeneration of nutrients occurs in water column
productivity is ultimately limited by regeneration of
nutrients from deeper waters
(c) 2001 by W. H. Freeman and
Company