SECONDARY PRODUCTION

THIS BASICALLY MEANS THAT:

INTRODUCTION:  R (RESPIRATION RATE) - depends on temperature
Within any given ecosystem, primary production refers to the  EI - energy needed for the process (in this case,
processes by which autotrophs such as plants, use sunlight to RESPIRATION).
produce chemical energy, which can be passed to other  K - Boltzmann constant (a physical constant used in
consumers including animals, fungi and heterotrophic bacteria. thermodynamics)
Consumers serve two principal needs: they metabolize the  T – temperature (Kelvin).
organic substrate wasting energy and renew basic sources for
productivity of the Qualitative Dimensions, secondary - As temperature increases, respiration rates correspondingly
production, and apply part of this matter for their own need, rise, due to accelerated metabolic processes.
which gives rise to secondary production, which can be pushed
higher up the food chain. METABOLIC RATES VS TEMPERATURE

The energy the rabbit gets from food (like the carrot in the
diagram) is broken down into different parts

This graph demonstrates that:

1. Ingestion – is the energy the rabbit takes in when it eats
 Metabolic rates positively correlate with temperature.
food
2. Assimilation – part of the ingested energy that the  Homeotherms (warm-blooded) showcase higher
rabbit’s body absorbs and uses metabolic rates compared to poikilotherms (cold-
3. Egestion – energy that is lost because the rabbit blooded).
cannot digest all the food
4. Respiration – Some energy is used for respiration METABOLIC RATES VS BODY MASS
(breathing and other body processes). This energy is
lost as heat.
5. Growth or Production – Energy used for growth or
reproduction (like growing bigger or having babies)
6. Excretion – energy that is lost through urine (waste).

Energy Budget Equation

 Energy taken (I) = energy absorbed (A) + Energy lost
in feces (E)

 Energy absorbed (A) is split into energy lost through

respiration (R), + used for growth (P) or lost in urine
(U). Observations indicate that smaller organisms tend to exhibit
higher metabolic rates per kilogram of body mass than larger
organisms.

The Formula they give is

METABOLIC RATES VS BODY MASS EFFICIENCY VARIABILITY BY ORGANISM TYPE
Typical efficiency values for various organisms are summarized:

This graph indicates a trend: smaller animals utilize more

oxygen per gram than larger ones, implying higher energy
demands relative to size.
Mathematical Constants in Energetics SECONDARY PRODUCTION
The chapter on secondary production emphasizes its crucial
role in the ecological energy budget for two reasons:
 Relative ease in estimating secondary production
compared to assimilation and respiration.
Key definitions include:  An inclination towards viewing food webs as energy
 e: Base of natural logarithms, approximately 2.718, transfer mechanisms enhancing human sustenance.
relevant for growth and decay calculations. Common misconceptions exist regarding:
 EI: Activation energy required for biochemical  Confusion between secondary production and
reactions. biomass accumulation.
 k: Boltzmann's constant, relating the energy of  Misinterpretation of gross versus net secondary
particles to temperature. production.
Q10 Temperature Dependence Factor Government population stability despite fluctuation in births and
deaths highlights the complexities of estimating biomass
 The Q10 factor expresses the rate of increase in accumulation.
metabolic processes with each 10°C rise in
temperature, generally implying that rates tend to Method for Estimating Secondary Production
double.
Secondary production has been studied through various
methods, focusing on:
ENERGY PARTITIONING AMONG CONSUMERS
 Biomass and growth rate analysis of different size
Animal energy utilization encompasses multiple efficiencies:
classes.
 Assimilation Efficiency (A/I): Reflects the rate at
 Body mass fluctuations within populations.
which energy is extracted from food.
 Mean annual biomass calculations and productivity
 Net Growth Efficiency (P/A): Compares energy
assessments.
directed towards growth against that expended on
metabolic processes.
 Gross Growth Efficiency (P/I): Represents the
conversion efficiency of consumed food into body
tissue.
where Bi and gi are the standing biomass and instantaneous PRODUCTION OF AN INDIVIDUAL SPECIES OF
growth rates of the ith size class, respectively. For exponential CONSUMER
growth,
The studies by Plante and Downing (1989), Morin and Bourassa
(1992), Benke (1993), Tumbiolo and Downing (1994), and
Cusson and Bourget (2005) examined the prediction of aquatic
invertebrate consumer production using empirical data. All
reached similar conclusions, highlighting consistent patterns in
the factors influencing production (see Figure 3.4).

Where: Di = the development time (the length of time the animal

spends in the ith size class) mb+1 and mb = the body masses
of the animals entering and leaving the ith size-class.

Where: N = the mean population density t = time mt and mt11 =

mean body masses at two successive sampling times

 Secondary production can be predicted with a 5-fold

confidence interval (25-fold total range) using
Where: B = mean annual biomass P/B = annual turnover ratio. average annual biomass, body size, and
temperature.
 Biomass is the strongest predictor of production;
adding body size and temperature to models provides
minimal improvement in prediction.
Controls and Predictability of Secondary Production
 Model coefficients:
Researchers have focused on understanding primary
production control factors, such as nitrogen, phosphorus, water,  `Production is linear with biomass (B^1.0).
and light, although less attention has been devoted to
secondary production:  Production is inversely proportional to the
fourth root of body mass (M^-0.25).
 Secondary production is studied at the individual
population level rather than as a collective  Production is temperature-dependent with a Q10
measurement. of 2 to 2.5.

 It proves least predictable at the individual species  These relationships align with the mass-
level, while more accurate predictions emerge at the dependence of metabolic processes (Peters
community level. 1983).

Key points include: PRODUCTION OF AN INDIVIDUAL SPECIES OF

CONSUMER
 Individual population production relies heavily on
biomass, temperature, and body size. Tumbiolo and Downing (1994) found that marine invertebrate
production was higher in shallow waters and seagrass beds,
 Even empirical models face challenges due to self- where food quality was better. This led to a higher production-
correlation and difficulties in biomass estimation. to-biomass ratio (P/B), indicating faster growth and mortality,
while deeper, food-poor waters showed slower life rates.
 Empirical models for predicting animal population
production often rely on mean annual biomass, but these
models come with significant error.
  Criticism by Benke (1993): These models are PRODUCTION OF THE ENTIRE COMMUNITY OF
statistically invalid due to self-correlation, as production CONSUMERS
is directly linked to biomass.
 Biomass as a predictor:

 Not ideal: Estimating mean annual biomass is costly

and prone to errors.
 While biomass data is essential, using it for predicting
production doesn't significantly improve the model's
value.
Unlike previous cases with statistical estimates, the total
Challenges with alternative models: production of a consumer community in an ecosystem can be
calculated exactly in principle (Strayer 1988). For example, in
 Models without biomass: These are
an ecosystem receiving 100 g of organic matter and populated
underdeveloped and likely to be highly error-prone.
by consumers with a 30% net growth efficiency, the total
Consumer density (N): production can be determined directly.
 A key factor in predicting production, but varies widely In this example, an ecosystem receives 100 g of organic
and is currently unpredictable due to numerous matter. After the first round of consumption, 70 g is lost to
ecological factors. respiration, and 30 g remains in the system. In the second round
of consumption, 30 g is processed, producing 9 g of production
 Existing density models (e.g., Peters 1983) are and 21 g of respiration. In the third round, 9 g is consumed,
imprecise, making it difficult to predict production resulting in 2.7 g of production and 6.3 g of respiration. By
without relying on biomass. the time all of the original 100 g is respired, a total of 54.2 g of
Production of an guild of a Consumers secondary production will have occurred.

Studies of both animals and bacteria (Kajak et al. 1980; Cole et Conclusions on Predicting Secondary Production
al. 1988; McNaughton et al. 1989, 1991; Wallace et al. 1999; cf. Several conclusions emerge regarding secondary production
also Cyr and Pace 1993; Cebrian 1999) have shown that the predictions:
production of a functional group of consumers is correlated with
the supply rate of its food (Figure 3.5). 1. Predictive accuracy improves at the community level
due to constraints governing food supply and biomass.
2. Organic matter supply and retention processes among
consumers exert significant influence.
3. Further research is necessary to explore the
relationships between nutrient availability, organic
matter transfers, and community dynamics.

ORGANIC MATTER DECOMPOSITION

DECOMPOSITION
Decomposition refers to the breakdown of organic matter into
simpler forms. This process occurs at two scales:
 Macroscale: Involves the conversion of large,
recognizable materials into smaller particles, soluble
compounds, and gases. Here, organic materials lost
from the parent material can serve as food resources
for other organisms. Fine particles and soluble
compounds can be transported across ecosystems,
influencing food webs.
 Molecular Scale: Complex organic molecules like
carbohydrates and proteins are transformed into
simpler forms, such as sugars and amino acids,
 ultimately leading to inorganic compounds like NET GROWTH EFFICIENCY (NGE)
ammonium and carbon dioxide.
NGE is calculated as follows:

Where G represents growth and R represents respiration. NGE

describes the proportion of assimilated carbon available for
consumption by heterotrophs.

MICROBIAL PROCESSES IN DECOMPOSITION

Microbial processes play a significant role in decomposition.
Microbes can:
 Enhance the digestibility of detritus through enzymatic
action.
 Contribute to the pool of assimilable carbon.
 Regenerate essential nutrients like nitrogen and
phosphorus, making them available for plants and
microbes, which is crucial for ecosystem productivity.
The availability of limiting nutrients, such as nitrogen and
phosphorus, often controls ecosystem productivity,
underscoring the importance of decomposition rates.
COMPARING AQUATIC AND TERRESTRIAL
DECOMPOSITION
 Moisture Availability: Influences organic matter
ROLE OF INVERTEBRATES IN LITTER DECOMPOSITION
decay in terrestrial systems.
Invertebrates are adapted to ingest large litter particles, which
 Soil Formation: Processes like humification are more
increases mass loss in both aquatic and terrestrial ecosystems.
prominent in terrestrial ecosystems due to minimal loss
of decomposition by-products compared to aquatic  Shredders in Aquatic Ecosystems: These
systems, where physical transport can limit organisms break down leaf litter, which accelerates the
interactions. generation of small particles.
ORGANISMS RESPONSIBLE FOR DECOMPOSITION  Impact of Pesticides: Reduction of aquatic insect
populations, such as through pesticides in a North
Microorganisms, particularly bacteria and fungi, are responsible
Carolina stream, significantly decreased the export of
for the bulk of organic matter decay. Key points include:
fine particles.
 Bacteria: Have a turnover rate of 1 to 10 days.
Research Findings
 Fungi: Turnover rate varies from several days to
Hieber and Gessner (2002) found that macroinvertebrates and
weeks; fungal biomass is often more abundant than
microbes contribute equally to leaf mass loss using litter bags
bacterial biomass.
with varying mesh sizes.
 Fungal Structures: Most fungi have filamentous
Significance of Fine Particulate Organic Matter (FPOM)
structures (hyphae) that penetrate plant cell walls,
requiring sufficient hyphal mass for reproduction. FPOM produced by shredders is quantitatively significant and
serves as food for diverse filtering and collecting organisms.
 Functional Differences:
o Fungi excel at degrading tough materials.
o Bacteria are more efficient at acquiring
nutrients.
  First-order Model: The rate of mass loss is
proportional to the mass still present.
 Exponential Constant (K): A measure of differences
in rates across types of litter, temperature, and
presence/absence of certain organisms.
Example
Enriquez et al. (1993) reported median k values for various plant
groups ranging across three orders of magnitude:
 Perennial leaves from broadleaf trees – slowest.

ROLE OF DETRITUS IN MACROCONSUMER PRODUCTION  Seagrasses and macroalgae – intermediate.

A significant portion of secondary production in terrestrial and  Microalgae – fastest.

aquatic ecosystems is derived from detritus. MASS LOSS AND ORGANISMS
 Increases in microbial abundance and activity improve Some products of mass loss serve as food resources for other
the digestibility of detritus for macroconsumers. organisms. The mineralization of organic carbon to CO2
 Microbial biomass is the digestible portion, while much represents a dead end in terms of chemical potential energy.
detrital organic matter is less usable for consumers. Factors Affecting Mass Loss
PEANUT BUTTER AND CRACKERS MODEL: Loss of particles depends on three main factors:
This model illustrates the relationship between microbial and  The fragility of the original material.
plant material:
 Physical stresses in the system.
 Microbes: Represent the digestible "peanut butter".
 The prevalence of animals capable of breaking litter
 Plant Material: Represents lower quality, less into smaller pieces.
digestible "crackers".
Early Mass Loss Phase
MICROBIAL INFLUENCE ON DETRITUS
Loss of soluble organic compounds dominates the early mass
Microbial degradation influences the supply of carbon for loss phase of decomposition, leading to a rapid decline over the
consumers in significant ways: first few days, followed by slower rates and a flattening of the
 Microbes release extracellular enzymes that curve.
"predigest" plant macromolecules, enhancing DISSOLVED ORGANIC MATTER
digestibility.
Dissolved organic matter released from litter in aquatic systems
 Even with low assimilation efficiencies, large quantities or leached from soils can:
of ingested plant material contribute to carbon needs.
 Support food webs.
 High microbial degradation can reduce standing stocks
of detritus, depleting food resources for consumers.  Interact with inorganic nutrient pools.

MICROBIAL INFLUENCE ON DETRITAL ORGANIC MATTER  Alter light regimes.

Microbial biomass enhances the palatability and availability of  Complex metals and toxic organic compounds in
detrital organic matter for invertebrates, but microbial diverse ways (Findlay and Sisabaugh, 2003).
metabolism can also convert organic carbon into CO2, limiting
Ultimate Fate of POC
food availability for consumers.
The ultimate fate of particulate organic carbon (POC) is
Dissolved Organic Carbon (DOC) supports bacterial growth but
mineralization to CO2, which is of no further energetic use to
is typically unavailable as food for macroinvertebrates.
heterotrophs and often leaves the aquatic ecosystem.
Decay Rates Generation of methane (CH4) is susceptible to oxidation by
appropriate microbes (e.g., methane oxidizers), contributing to
The shape of the curve is often represented by a negative secondary production.
exponential model:
MEASUREMENT OF MASS LOSS IN LITTER phenolics and tannins, can resist breakdown and inhibit enzyme
activity, although their effects vary across species and
Measurement of mass loss from litter bags has been informative environments.
and provides a comparative and integrative approach to
studying decomposition. However, this method limits our EXTRINSIC FACTORS
understanding of specific processes responsible for mass loss
and the recycling of detrital organic matter. Factors like temperature and moisture also affect decay rates:

This technique also cannot study long-term decay processes,  Nutrient-rich environments enhance decomposition by
as gases, small particles, and soluble compounds do not remain boosting microbial biomass.
in the litter bag. The decay of these components is often studied  Low pH and oxygen absence reduce decay due to
under artificial conditions of containment or by measuring net limitations on enzyme activity and microbial efficiency.
CO2 released from fine sediment or soil.
 Increased species diversity in both litter and
CONTROLS ON DECOMPOSITION decomposers generally enhances decay rates, though
Decay Rates mixing different plant litter species produces varied
results.
 Intrinsic Factors: Plant-specific traits that influence
decay rates include plant nitrogen content, biochemical INTERACTIONS WITH ELEMENT CYCLES
composition, and litter quality. Key elements lacking, such as nitrogen and phosphorus,
 Extrinsic Factors: Environmental factors such as contribute to nutrient shortages in plant litter. Many plant litters
nutrient availability, herbivores, moisture, and are characterized by a relative shortage of essential nutrients,
temperature also play crucial roles in decomposition particularly nitrogen and phosphorus, which can challenge
rates. microbial decomposition.

The interaction between intrinsic and extrinsic factors can shape MICROBIAL ROLE
decomposition rates significantly. For example: Microbes play a vital role by:
 Nutrient-poor environments might cause plants to  Utilizing detrital materials as carbon sources.
evolve higher concentrations of structural
carbohydrates or lignin, making them harder to  Assimilating inorganic nutrients from the environment.
decompose.
IMPACT OF RISING CO2
 Herbivore activity can alter plant nitrogen content by
The enrichment of plant tissues in carbon leads to an increased
inducing plant defense mechanisms, indirectly
need for nitrogen and phosphorus for effective decomposition,
affecting decomposition rates.
highlighting the influence of human-induced changes on these
LIGNIN processes.
Lignin is a complex macromolecule composed of diverse CONCLUSION
subunits containing phenolic rings. Due to its complexity:
Understanding decomposition processes, the role of
 Lignin is difficult to break down, resulting in decay microorganisms, and the intricate relationships with nutrient
resistance and lower energy content. cycling is crucial for addressing ecological concerns and
enhancing ecosystem productivity.
 The higher the lignin content in plant material, the
slower the decomposition rates.
STRUCTURAL MACROMOLECULES
The physical resistance of lignin and lignocellulose provides
structural support to plants, making them resistant to decay.
Extracellular enzymes are required for the breakdown of these
polymeric compounds.
NUTRIENT IMPACT ON DECOMPOSITION
Nitrogen and phosphorus are often scarce in plant litter, limiting
microbial decomposition. Low nitrogen content restricts
microbial growth and carbon metabolism, leading to incomplete
decomposition. Secondary plant compounds, including
ELEMENT CYCLING o Soil weathering, facilitating absorption by
plants.
Element cycling refers to the movement and transformation of
chemicals within and between ecosystems, serving as a o Transfer of elements through various trophic
crucial focus for ecosystem scientists. It connects living levels within biotic communities.
organisms with nonliving components, highlighting the
importance of nutrient cycles that include critical elements  Linked Elemental Cycles
such as nitrogen, phosphorus, potassium, and calcium The biotic and abiotic spheres consist of diverse elements:
essential for plant growth and ecosystem productivity.
o Over 20 elements are crucial for plant
growth and reproduction.
Introduction to Element Cycling o Minerals contain a combination of elements,
Nutrient cycling plays a vital role in various aspects of primarily oxygen, silicon, aluminum, and iron.
ecosystem science, including primary production, food webs, Categories of Linkages
and decomposition processes. Addressing environmental
issues often requires managing imbalances in elemental o Structural Stoichiometry
availability vital for human food production.
 Involves the ratio of elements
 Macronutrients: present in materials or organisms.
o Nitrogen and phosphorus receive significant o Functions of Chemical Reactivity
attention due to their importance in
 Concerns how chemical changes
ecosystem productivity.
influence material and energy flow
 Trace Elements: through ecosystems.
o Elements like copper, although essential in o Stoichiometry in Ecosystems
trace amounts, can become toxic at higher
Ecological stoichiometry studies how elemental proportions
concentrations, demonstrating their dual role
within organisms and their environments influence biological
in pest and algae management.
processes and nutrient cycling.
What is an Element Cycle?
Key Contributions
An element cycle encompasses the various pathways through
 Redfield Stoichiometry: Alfred C.
which both organic and inorganic materials move. These
Redfield (1958) found that marine
pathways can be characterized as transformation processes
phytoplankton typically contain
between:
carbon, nitrogen, and phosphorus
 Abiotic and Biotic Pools in a ratio of 106:16:1, which mirrors
those in ocean water.
 Within Biotic Pools
 Sterner and Elser (2002):
 Within Abiotic Pools Explored stoichiometric
 Physical Movement Between Pools requirements of primary producers
and consumers, linking these needs
In general, materials flow between these pools as they are to ecological adaptations.
absorbed by organisms (production) and subsequently broken
down (decomposition). Redox Reactions

Movement and Transformation of Elements Central to elemental cycles, redox reactions facilitate the
transformation and movement of elements among various
 Elements are integrated into living biomass through chemical forms in the environment.
uptake by organisms.
The Importance of Chemical Properties
 Upon organism death, the decomposers break down
organic matter, releasing elements for reuse by other Chemical properties—determined by an element’s placement
organisms. in the Periodic Table—impact nutrient cycling and ecosystem
dynamics. Elements may:
Inorganic and Organic Material Pathways
 Form cations by losing electrons.
 Movement occurs through:
  Gain electrons to form anions. Example: The Sulfur Cycle
 Remain inert, such as noble gases. In a northern hardwood ecosystem, sulfur enters through
atmospheric deposition and interacts with other elements while
Oxidation States moving through the ecosystem. Sulfur uptake by plants
Key elements (carbon, nitrogen, sulfur, oxygen) exhibit multiple exemplifies how sticking and chemical transformation interplay
oxidation states, engaging in redox reactions that facilitate in elemental cycling.
cycling through different forms. Questions and Research Directions
Nutrient Spiraling Ongoing inquiries in ecosystem science focus on:
Nutrient spiraling outlines the transportation and recycling of  The effects of biotic and abiotic
nutrients within ecosystems, creating a spiral pattern from factors on nutrient release.
streams to sediments.
 Considering the implications of
Main Parameters these dynamics for ecosystem
 Areal Uptake (U): The dissolved management.
nutrient removal rate from water per Future chapters will elaborate on the cycling of carbon,
area over time. nitrogen, and phosphorus while prompting explorations into
 Uptake Length (Sw): The distance other elemental roles in ecosystem processes.
nutrients travel in water before
uptake.
 Uptake Velocity (vf): The rate of
nutrient transfer from water column
to sediment.

Findings on Nutrient Dynamics

Research indicates ammonium (NH₄⁺) uptake occurs faster
than nitrate (NO₃⁻) uptake, especially in low-discharge
streams. Conversely, higher discharge streams see extended
nutrient uptake lengths.
Conclusion
A comprehensive understanding of chemical properties and
nutrient spiraling informs effective management of aquatic
ecosystems, equipping ecologists with tools for monitoring
stream health.
Framework for Elemental Cycling
The fate and movement of materials within ecosystems can be
encapsulated in a three-part framework:
9. Movement: Material migration within systems
influenced by physical, chemical, or biological vectors.
10. Sticking: Temporary retention of elements within
systems, crucial for reducing losses over varying durations.
11. Change: Transformation of materials among
chemical forms, influencing their travel or retention and energy
dynamics.