HOW A RAINFOREST

FUNCTIONS

Dawn R. Black
 Questions
1. What factors influence productivity?
2. How does primary productivity in tropical
rainforests compare to other biomes?
3. Where are most of the rapidly recycling
minerals in tropical rainforests found?
4. What are the three general types of soils
found in the tropics?
 Questions
5. What are the nutrient retention adaptations
found in oligotrophic soils?
6. How do rainforest plants receive nitrogen?
 GPP & NPP/Biomass
Gross Primary Productivity (GPP)
The total amount of photosynthesis accomplished
Net Primary Productivity (NPP)
Amount of carbon added to the plant for growth and
reproduction
Biomass + Detritus + Soil Organic Matter
Biomass
Total storage of organic carbon in plant tissues
Factors Influencing Productivity
Adequate light (low light intensity limits
understory species)
Moisture
CO2 levels
Soil minerals/nutrients (many soils old and
mineral poor)
 Tropical vs. Other Ecosystems
GPP vastly higher in rainforests than in any
other ecosystem
High rates of respiration (temperature
stress)
50-60% of GPP spent on maintenance
NPP higher than any terrestrial ecosystem
 Comparisons of NPP
Net Primary Production (NPP) of the Major Biome Types Based on Biomass Harvestsa.

Aboveground Belowground NPP Belowground Total NPP

NPP (g m-2 yr-1) NPP (g m-2 yr-1)
Biome (g m-2 yr-1) (% of total)
Tropical forests 1,400 1,100 0.44 2,500
Temperate forests 950 600 0.39 1,550
Boreal forests 230 150 0.39 380
Mediterranean shrublands 500 500 0.50 1,000
Tropical savannas and 540
grasslands 540 0.50 1,080
Temperate grasslands 250 500 0.67 750
Deserts 150 100 0.40 250
Arctic tundra 80 100 0.57 180
Crops 530 80 0.13 610
a
Data from Saugier et al. (2001). NPP is expressed in units of dry mass. NPP estimated
from harvests excludes NPP that is not available to harvest, due to consumption by
herbivores, root exudation, transfer to mycorrhizae, and volatile emissions.
Global Distribution of Carbon in
Plant Biomass
Global distribution of terrestrial biomes and their total carbon in plant biomassa.

Area (106 km2) Total C pool Total NPP

(Pg C) (Pg C yr-1)
Biome
Tropical forests 17.5 340 21.9
Temperate forests 10.4 139 8.1
Boreal forests 13.7 57 2.6
Mediterranean shrublands 2.8 17 1.4
Tropical savannas and grasslands 27.6 79 14.9
Temperate grasslands 15.0 6 5.6
Deserts 27.7 10 3.5
Arctic tundra 5.6 2 0.5
Crops 13.5 4 4.1
Ice 15.5
Total 149.3 652 62.6
a
Data from Saugier et al. (2001). Biomass is expressed in units of carbon, assuming that
plant biomass is 50% carbon.
 Productivity by Biome
Productivity per day and per unit leaf areaa.

Season Daily NPP per Daily NPP per

lengthb ground area Total LAI leaf area (g m-
c

Biome (days) (g m-2 d-1) (m2 m-2) 2 -1

d )
Tropical forests 365 6.8 6.0 1.14
Temperate forests 250 6.2 6.0 1.03
Boreal forests 150 2.5 3.5 0.72
Mediterranean shrublands 200 5.0 2.0 2.50
Tropical savannas and grasslands 200 5.4 5.0 1.08
Temperate grasslands 150 5.0 3.5 1.43
Deserts 100 2.5 1.0 2.50
Arctic tundra 100 1.8 1.0 1.80
Crops 200 3.1 4.0 0.76
a
Calculated from Table 5.3. NPP is expressed in units of dry mass.
b
Estimated
c
Data from Gower (In press).
 Tropical vs. Temperate PP
Huston (1994)
Productivity per unit time no greater in the tropics than
in temperate zone (high PP due to length of growing
season)
Kricher (1997)
Maybe plant tissue grows faster in tropics
Tropical species grew by an order of magnitude more
than temperate species (red oak, red maple) when
length of growing season was corrected for
Suggests that per tree productivity is considerably
enhanced in the tropics
 Nutrient Cycling

Decomposition and subsequent recycling is

the process by which materials move
between the living and nonliving
components of an ecosystem
 Decomposition
Fungi & bacteria convert dead
organic tissue back into simple
inorganic compounds reavailable
to plant root systems
Fungi immensely abundant in tropics
Mycelial mesh covers parts of some
tropical forest floors
 Supporting Decomposers
Slime molds
Actinomycetes
Algae
Animals (vultures, arthropods, earthworms,
invertebrates)
Protozoans
 Leaching of Nutrients
Leaching washing of essential minerals
and other chemicals from leaves and soils
by water
 Leaf adaptations

Drip tips (speed water runoff)

Protective cuticle with lipid-soluble

secondary compounds that retard water loss
& discourage herbivores and fungi
 Leaching of Soil
Rainfall increases H+ ions in soil (lowers
pH), which bind to (-)-charged humus &
clay
(+)-charged minerals (Ca, K) washed to
deeper part of soil
Acidity of soil increased
 Rapid Recycling of Nutrients
Most of rapidly recycling minerals are in
the biomass in the tropics
Decomposition & recycling of fallen parts
occur with much greater speed in rainforests
than in temperate forests (thin litter layer)
~80% of total leaf matter in Amazon rainforest
annually returned to soil (Klinge et al. 1975).
 Role of Mycorrhizae
Substitute for poorly developed root hairs
Mostly vesicular-arbuscular (VAM)
Aid in uptake of phosphorous

Some ectomycorrhizae, especially in poor soils

Aid in uptake of both minerals and water

VAM status of Dicorynia guianensis seedlings is

critical factor controlling regeneration in primary
tropical forest of French Guiana (Bereau et al.,
1997)
 Soil Characteristics
Determined by several factors (Jenny 1941):
Climate
Vegetation
Topographic position
Parent material
Soil age
 Rainforest Soil Types
Three general classifications of soils throughout
humid tropics
1. Ultisols
2. Oxisols
3. Alfisols

Comprise ~71% of land surface in humid tropics

worldwide
Only ~15% of moist tropical forests moderately fertile
(in young soils of recent origin)
 Ultisols
Well-weathered

Minerals leached from

upper parts of soils
 Oxisols
Deeply weathered
Old
Acidic
Found on well-drained
1M
soils of humid regions
Also found on Guianan
Shield (common
throughout global tropics)
Reddish color due to iron
& aluminum oxides
 Alfisols
Closer to neutral pH
(still acidic)
Less overall
leaching
Common in
subhumid &
semiarid tropics
 Mineral Cycling on Oligotrophic Soils

Up to 26% of roots on the surface

Root mats several cm thick can develop
Root mat & mycorrhizae directly absorb available
minerals
99.9% of Ca & P absorbed into root mat in
Amazon
Presence of buttresses may allow roots to spread
widely at surface, where they reclaim minerals
 Nutrient Retention Adaptations
Surface roots/mats
Apogeotropic roots roots grow upward from soil
onto stems of neighboring trees, absorb nutrients
leached from trees from throughfall
Arrested litter epiphytes & understory plants
catch litter from canopy
Canopy leaves algae & lichens on leaves absorb
nutrients from rainfall and trap on leaf
 Nitrogen Fixation
Legumes & Rhizobium abundant in biomass &
biodiversity in tropics, take up gaseous N from
atmosphere & convert to nitrate
Certain epiphytic lichens fix nitrogen
Leaf-surface microbes & liverworts may facilitate
uptake of gaseous nitrogen
Termites N-fixation due to activities
of microbes in termite guts
 Rainforest Gaps
Microclimates dependent on gap size
Affects light, moisture, & wind conditions
Treefalls are normal part of rainforest
function, peak in rainy season
Creates heterogeneous forest
 Gap-Dependent Pioneer Species
Produce an abundance of small seeds
dispersed by bats or birds
Seeds capable of long dormancy periods
Different growth patterns among pioneers
may explain coexistence of so many
different species in rainforest ecosystems
 Forest Demographics
Forest turnover varies with species & region
La Selva, Costa Rica ~118 years
Cocha Cashu, Peru 63 years
Manaus, Brazil 82-89 years
 Disturbance & Ecological
Succession
Jungle = early succession in tropics
High species richness
Highly variable from site to site
Early succession Colonizers
Small in stature, grow fast, produce many-seeded fruits
Late succession Equilibrium species
Larger, grow more slowly, fewer seeds per fruit, persist
in closed canopy
Can take >500 years to reach equilibrium
 Answers
1. What factors influence productivity?
Light levels, moisture, CO2 levels, soil
minerals/nutrients

2. How does primary productivity in tropical

rainforests compare to other biomes?
Both GPP & NPP are higher than other
biomes
 Answers (cont.)
3. Where are most of the rapidly recycling minerals
in tropical rainforests found?

In the plant biomass

4. What are the three general types of soils found in

the tropics?

Ultisols, Oxisols, Alfisols

 Answers (cont.)
5. What are the nutrient retention adaptations
found in oligotrophic soils?

Surface roots/mats, apogeotropic roots,

arrested litter, algae/lichens on leaves
6. How do rainforest plants receive nitrogen?
Legumes & Rhizobium, epiphytic lichens,
leaf-surface microbes/liverworts, termites
 Roggy et al. (1999)

Study of plant N nutrition in legumes &

pioneer species at Piste de St Elie in the
ECEREX research area, French Guiana
Used 15N method to estimate nitrogen
input by N2-fixing legumes to natural
rainforest
 Roggy et al. (1999)

Results
N2-fixing legumes contributed 136 t ha-1 to total
above-ground plant biomass
N2-fixation estimated to be 7 kg ha-1 y -1
15N of non- N2-fixing plants could be related
to soil nitrogen availability
Could be used as indicator of nitrogen-cycling
efficiency in rain forests
 Chave et al. (2001)
Biomass study
2 study sites
Nouragues Research Station (100 km
inland)
Piste de Saint-Elie Research Station
(coastal rain forest
 Chave et al. (2001)
Results
Significant spatial variability of biomass at
fine-scale resolution
Illustration of disturbance-driven, mosaic-like
pattern in old-growth forest
Biomass accumulation of 3.2 Mg ha-1 y -1 &
2.8 Mg ha-1 y -1, which agrees with literature
NPP of 2-4 Mg ha-1 y 1 (Phillips et al., 1998)
-Variability of biomass correlated with canopy
gap openings
 Granier et al. (1996)
Transpiration of natural rainforest & its
dependence on climatic factors
Objectives:
Analyze transpiration at tree level through sap
flow measurements performed on several major
species growing in their natural environment
At stand level, analyze dependence of
transpiration to climatic factors, by scaling up
allowing calculation of stand.
 Granier et al. (1996)
Dependent Factors
Late stage species (high flow rates)
Pioneer species (low flow rates)
Crown Status
Codominant trees exhibited lower flow rates
than dominant trees of same species
Sap flow showed remarkable concordance
with variations of air vapor pressure deficit
 Literature Cited
Bazzaz, F.A. 1984. Dynamics of wet tropical forests and their species strategies. In E. Medina, H.A. Mooney, and C.
Vazquez-Yanes (eds). Physiological ecology of plants of the wet tropics. Junk, Dordrecht, pp. 233-243.
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seedlings of Dicoryniaguianensis Amshoff, a tree species of the tropical rain forest in French Guiana.
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