Secondary production is the generation of new heterotrophic biomass and is analogous to net primary production of autotrophs. For an individual, secondary production is equivalent to the growth of new somatic or reproductive biomass over time. For a population, secondary production comprises the total formation of biomass, regardless of its fate, by all individuals within the population over a defined time interval. Some consider secondary production the ultimate measure of population ‘success’ because it incorporates aspects of survivorship, individual growth rate, biomass, development time, and reproduction. Secondary production is often associated with the subfield of ecosystem ecology because it is a flux with dimensions of mass or energy area-2 time-1. This flux is typically estimated with an ecological currency (e.g., joules, carbon, organic matter) that can be compared with other ecosystem processes such as primary production or decomposition. Secondary production estimates are thus useful for placing species, populations, and communities within a broader ecosystem context, and for facilitating the study of energy flows and ecological efficiencies in trophic interactions. The vast majority of secondary production estimates have come from freshwater and marine ecosystems, while there are very few studies in terrestrial ecosystems. In the aquatic studies, although early work was largely focused on fishes, most estimates are for benthic invertebrates; some studies have quantified production of zooplankton, bacteria, and fungi. Early studies of secondary production were focused on methodology and basic comparisons among populations or communities. More recent literature has expanded the application of secondary production toward broader ecological questions related to, for example, energy and chemical flows in food webs, species interaction strengths, and responses to anthropogenic stressors. This bibliography focuses on primary literature that highlights key historic, conceptual, theoretical, and applied papers related to secondary production. Papers highlighted herein are biased toward freshwaters and invertebrates because of their dominance in the literature, but key references that extend to other habitats and taxa are included.
History and Context
Benke, et al. 1988 emphasizes that the measurement and application of secondary production has deep roots in bioenergetics, a topic which has been at the forefront of ecology since the mid-20th century. From a bioenergetics perspective, secondary production represents the energy assimilated by an individual or population that is not lost to respiration and is therefore available to other trophic levels. Some of the earliest estimates of secondary production in Lindeman 1942; Clarke, et al. 1946; and Odum 1957 were used to construct energy budgets for entire ecosystems. Although these estimates were relatively crude, and often lumped taxa into broad trophic categories, their application helped promote subsequent development and refinement of methodology. Secondary production is also a central theme in Odum and Odum 1953 in its conceptualization of energy flow in ecosystems and the application of thermodynamics to trophic interactions. Broad-scale measurement of secondary production was catalyzed in the late 1960s by the International Biological Programme, in which standardized methods were developed and applied among different taxa and ecosystems globally (see Ricker 1968, Edmondson and Winberg 1971, Holme and McIntyre 1971, and Sorokin and Kadota 1972). Benke 2010 shows that there are now thousands of secondary production estimates, with renewed emphasis on understanding its drivers.
Benke, A. C. 2010. Secondary production as part of bioenergetic theory—contributions from freshwater benthic science. In Special issue: Freshwater benthic science—What has it contributed to ecological theory? Edited by Alexander M. Milner and Klement Tockner. River Research and Applications 26.1:36–44.
This paper provides additional context on the relationship between secondary production and bioenergetics, as well as a review of the distribution of studies among different taxa and habitats.
Benke, A. C., C. A. Hall, C. P. Hawkins, et al. 1988. Bioenergetic considerations in the analysis of stream ecosystems. Journal of the North American Benthological Society 7.4:480–502.
This review synthesizes information about bioenergetics in stream ecosystems and provides a general and informative discussion of the connection between bioenergetics and secondary production.
Clarke, G. L., W. T. Edmondson, and W. E. Ricker. 1946. Dynamics of production in a marine area. Ecological Monographs 16.4:321–337.
One of the first studies to measure and describe secondary production for a marine ecosystem.
Edmondson, W. T., and G. G. Winberg. 1971. A manual on methods for the assessment of secondary productivity in fresh waters. IBP Handbook No. 17. Oxford, UK: Blackwell Scientific Publications.
A manual for estimating secondary production of freshwater organisms, including zooplankton and benthos. Chapters also deal with sampling, insect emergence, feeding, assimilation, and respiration. This manual was published as part of the International Biological Programme.
Holme, N. A., and A. D. McIntyre. 1971. Methods for the study of marine benthos. IBP Handbook No. 16. Oxford: Wiley Blackwell, UK.
This manual on methods for marine benthic production was published as part of the International Biological Programme.
Lindeman, R. L. 1942. The trophic-dynamic aspect of ecology. Ecology 23.4:399–417.
A classic work that quantifies an energetic food web in a freshwater lake. This paper introduces and operationalizes the concepts of productivity, energy flow, trophic efficiencies, and succession.
Odum, H. T. 1957. Trophic structure and productivity of Silver Springs, Florida. Ecological Monographs 27:55–112.
Another classic that is considered one of the first complete energy budgets for an entire ecosystem—in this case, a large freshwater spring.
Odum, E., and H. T. Odum. 1953. Fundamentals of ecology. Philadelphia: B. Saunders Co.
Considered the first classical ecology textbook. Provides a basic explanation of principles and concepts related to energy in ecological systems.
Ricker, W. E. 1968. Methods for assessment of fish production in fresh waters. IBP Handbook No. 3. Oxford, UK: Blackwell Scientific Publications.
This manual provides methods for measuring fish production and was published as part of the International Biological Programme.
Sorokin, Y. I., and H. Kadota. 1972. Techniques for the assessment of microbial production and decomposition in freshwaters. IBP Handbook No. 23. Oxford, UK: Blackwell Scientific Publications.
This manual on methods for microbial production was published as part of the International Biological Programme.
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