Chapter 1. Assessment of Secondary
Production: the First Step
JOHN
A.
DOWNING
1 Introduction
This manual is designed to help freshwater ecologists choose methods for use
in the scientific study of secondary productivity. Secondary production has
been defined many times in the literature (e.g. Clarke 1946; Ivlev 1966; Allen
1971: Winberg 1971a,b: Waters & Crawford
1973; Edmondson
1974;
Cushman et al. 1978; Benke & Wallace 1980) and most definitions are in
agreement. Waters & Crawford (1973) use the term in the sense of Clarke
(1946) as 'that amount of tissue elaborated per unit time per unit area,
regardless of its fate'. Other definitions stress that reproductive products and
production lost to predators and other losses must be included. The tissue
elaboration that is usually considered to be 'secondary production' is the
production not only of herbivores but of all freshwater invertebrates (see
Morgan et al. 1980). Therefore, the rate of secondary production can be
defined more specifically for this manual as that amount of tissue elaborated
by freshwater invertebrates per unit time per unit area, regardless of its fate
(after Clarke 1946; Waters & Crawford 1973). Many techniques exist for the
study of secondary production in freshwaters, and it is the goal of this book
to help the researcher to choose the appropriate ones to use under different
circumstances.
Although each author contributing to this handbook has dealt with a
different
set
of techniques,
one
single
conclusion
has
been
reached
independently by each. This common conclusion is that the choice of proper
technique depends upon the question posed or the hypothesis under
examination.
Many of the authors have come to a worrying second
conclusion. They believe that few production biologists to date have posed
questions or tested hypotheses; most have simply concerned themselves with
the estimation of single rates of production or its components. Because the
choice of technique depends upon the hypothesis to be tested, it has been
difficult for production ecologists to choose among the many techniques
available. The gravity of this conclusion has been discussed by many
philosophers of science. For example, F.S.C.Northrop (1947) has written
that 'One may have the most rigorous of methods during the later stages of
I
2
Chapter I
investigation, but if a false or superficial beginning has been made, rigor
later on will never retrieve the situation.' When questions are only posed
a posteriori, we risk the frequent choice of inappropriate methods (cf. LeCren
1972).
Because of this problem, this first chapter will review the general reasons
why ecologists estimate secondary production, and will then provide a
summary of the many interesting hypotheses suggested by the rich literature in
this field.
2 Theoretical Justification for Secondary Production Research
A field of study is usually judged useful if it has a potential for contributing to
established disciplines or goals. It is the same for the field of secondary
production
in
freshwaters.
Although
some production
biologists have
estimated productivity of a species in a certain area merely because no such
data have been published, many others feel that their studies are important
because they address one or more of four main conceptual subject areas.
These are:
(1)
The elucidation of energy or material transfers within communities and
(2)
The rational management of aquatic resources.
(3)
The detection of the effects of pollution.
(4)
The formation of general theories of biological productivity.
ecosystems.
Below, I present a brief discussion of the relationship between production
biology and these general ecological goals.
2.1 Energy or material transfer within ecosystems
G.E.Hutchinson (1942) has written that when Lindeman published his
famous paper The Trophic-Dynamic Aspect of Ecology' (1942), he hoped
that it would serve as a program for future ecological research. This has
certainly been true.
Lindeman suggested that if one could reduce the
interactions among components of a community to a common currency (e.g.
energy), then one could quantify the interactions and learn to predict changes
such as succession within ecosystems.
Lindeman
introduced the major
concept that an organism's success in an environment might be a function of
its ability to fix and retain energy.
This concept not only underlies much of current productivity research, but
was part of the stimulus for the research undertaken in the International
Biological Programme of which this handbook is a result. The elegance of this
concept is demonstrated by the frequency with which it has been accepted as
justification for research in secondary production (e.g. Kimerle & Anderson
Assessment of Secondary Production: The First Step
3
1971; Czeczuga & Bobiatyhska-Ksok 1972; Burke & Mann 1974; Nichols
1975; Zwick 1975; Benke 1976; Hibbert 1976; Zytkowicz 1976; Waters 1977;
Neves 1979; Benke & Wallace 1980; Tonolli 1980). I believe that Edmondson
(1974) has expressed it best: i cannot think of secondary production
as a distinct process by itself. Rather it is part of a larger scheme of
the movement of material through the ecosystem, and this is based on the
activities of individuals and populations of animals.' Much effort has gone
into the quantification of the components of this larger scheme (e.g. Kajak &
Hillbricht-Ilkowska 1972). The frustrating aspects are that even the simplest
community has many components, there are many different types of possible
interactions
among
components,
almost
all
individual
organisms
are
behaviorally plastic, and it is difficult to obtain accurate estimates of even one
rate of transfer under one set of simple circumstances. The result is that
fulfillment of the trophic-dynamic goal of production ecology is a formidable
task.
2.2 Management of aquatic resources
The measurement of secondary production is thought essential to the
management of aquatic resources, probably due to our trophic-dynamic view
of ecology. The most concrete freshwater resource is, of course, fish. Because
many fish depend to a high degree upon zooplankton and benthos for food
(e.g. Zelinka 1977), a variety of authors have suggested that an understanding
of the production processes of invertebrates will facilitate management offish
stocks (Zytkowicz 1976; Waters 1977; Williams c/ al. 1977; Priymachenko et
al. 1978) or prediction of rates offish production (Johnson & Brinkhurst 1971;
Moskalenko 1971; Czeczuga & Bobiatyhska-Ksok 1972; Johnson 1974;
Zytkowicz 1976). A recent paper by Hanson & Leggett (1982) shows that fish
yield is related to the mean standing biomass of macrobenthos in a lake, and
thus suggests that a general relationship probably exists between secondary
productivity and fish production. This relationship has yet to be described
empirically, however. The importance of secondary producers to the study of
fish dynamics (Hamill et al. 1979) is underscored by their trophic intermediacy
between fish populations and energy sources (Mathias 1971; Dermott et al.
1977). Johnson (1974) has also suggested that enhancement of secondary
production may be important to the development of freshwater aquaculture.
2.3 Detection of pollution
Because secondary production is a complex process that can be altered by
variations in many variables, it seems logical that variations in rates of
secondary production could be used to detect pollution (Winberg 1971b; see
review by Waters 1977). For example, Golterman (1972) found that the ratio
4
Chapter 1
of production to biomass (P/B) of zooplankton is higher in thermally polluted
waters than in control areas. A similar effect is suggested by McNaught&
Fenlon (1972). Many researchers have found that benthos production in lakes
is highest near areas of human activity (e.g. Mikulski et at. 1975;
Wolnomiejski et a/. 1976; Dermott et al. 1977). Zelinka (1977), on the other
hand, found that human activities (stream bed modification, toxic wastes, etc.)
most often have a negative effect on mayfly production. Other authors suggest
that secondary producers could be used in sewage treatment (e.g. Kimerle &
Anderson 1971; Waters 1977), or in the self-purification of polluted
ecosystems.
2.4 Formation of general theories of biological production
Winberg(1971a;Tonolli 1980) has stated that the'development of a theory of
biological productivity is one of the central aims of contemporary biology...'.
Mann (1972) has made a similar statement and suggests that we must 'make
every effort to improve the accuracy of the observations and the confidence
limits of resulting estimates' in order to help produce a general ecological
theory of biological budgets. If we take the term 'theory' in the usual sense,
that is, a construct that makes predictions about nature, then one of the basic
reasons for measuring secondary production is to learn how to predict it.
Looking back to Sections 2.1 and 2.2. we can see why it is very important to be
able to predict rates of productivity. The trophic dynamic analysis of
ecosystems requires the estimation of the secondary production of many
populations of animals. If these values could be predicted accurately under a
variety of conditions, then much effort could be saved in the trophic analysis
of communities. In addition, general theories of secondary production would
be very useful in the management of aquatic resources.
Brylinsky (1980) has written recently that productivity data should be
analyzed 'with a view to identifying those factors most important in
controlling biological production. Once identified, management efforts could
be directed towards manipulation of those factors appearing most important'.
Most of the balance of this introductory chapter will be devoted to an
exploration of those specific factors which have been suggested as important
in determining the rate of secondary production in fresh waters. It is my hope
that presentation of these hypotheses will help production biologists to define
specific questions for study, and thus indicate appropriate methods for
analysis.
3 Factors Affecting Rates of Secondary Production
This section contains a summary of the hypotheses suggested most frequently
by production biologists. Most of these hypotheses have arisen from isolated
Assessment of Secondary Production: The First Step
5
observations; only a few have been tested explicitly. It is not my intention to
suggest that these are the only interesting hypotheses or even the hypotheses
that will yield the most or quickest progress in production ecology. I only wish
to demonstrate that we possess a large set of implicit theories. These, or other
hypotheses, if tested explicitly, could not only yield progress in production
biology, but could make the choice of methods a more tractable problem.
For the sake of organization, I have arranged these hypotheses into four
categories. I will first discuss how rates of secondary production are affected
by characteristics of the population under study, then I will examine
hypotheses that relate to aspects of the environment in which they live.
Thirdly, I will present the few hypotheses in the literature that address the
manner in which secondary production is affected by interactions among
populations in the same community. Lastly, I will discuss the possible effects
of basin characteristics.
3.1 Effect ofpopulation characteristics
There are certain intrinsic characteristics of populations which dictate the
manner in which they live. When one examines an animal population casually,
certain elementary questions materialize. Four of these questions are: How
many are there, and what is their biomass? What is their life history like; how
long do they live, How big are they? What kind of animal are they?
Production biologists feel that each of these questions has a bearing on the
rate of secondary production that populations are able to attain.
3.1.1 Biomass
The literature contains a number of specific hypotheses regarding the
relationship of production (P) to mean biomass (B). First, there are many
(e.g. Laville 1971; Gak et ai 1972; Eckblad 1973; Waters & Crawford 1973;
Johnson 1974; Lavandier 1975; Mikulski et ai 1975; Wolnomiejski et al.
1976; Waters 1977; Hamill et al. 1979; Makarewicz & Likens 1979; Benke &
Wallace 1980; Short & Ward 1980) who have suggested that the ratio P/B is a
constant (c) for a given type of organism. That is:
P/B = c
(l.l)
If in fact P/B is constant, then production is an increasing linear function of
biomass with slope c and intercept zero:
P=cB
(1.2)
This relationship suggests that P = 0 at B =0, and that P = xc at B = oc. If
P/B is constant then production is not density dependent and is not subject to
6
Chapter 1
the normal constraints imposed by the carrying capacity of the environment.
A mental Malthusian exercise tells us that this cannot be so. Even though the
relationship between P and B may appear linear over a small range of B, the
convenient but inaccurate notion that P/B is constant should be abandoned.
Many have already done this for empirical reasons (e.g. McLaren 1969;
Schindler 1972; Paterson & Walker 1974; Jonasson 1975; Pedersen et al.
1976; Janicki & DeCosta 1977; Momot 1978; Pinel-Alloul 1978; Adcock
1979; Banse & Mosher 1980; Nauwerck et al. 1980; Uye 1982). Jonasson
(1975) has found that it is not even safe to use the same value of P/B for one
species at one site in successive years. He found that P/B for Chironomus
anthacinus was 4 in one year and 0-8 the next. Because the relation between
production and biomass is not linear, there will be a necessary negative
relationship between P/B and B. The danger is that variables correlated with B
(e.g. temperature, body size, respiration) may account for statistically
significant variation in P/B when they would not account for significant
variation in P beyond the accurately fitted effect of B. This could lead to errors
in both interpretation and predictive ability.
3.1.2 Age, lifespan, and voitinism
The length of life or relative age of individuals in a population also seems to
affect production. The influence of age on production is not clear-cut. Some
authors feel that P/B declines with age (Hibbert 1976; Waters 1977; Banse &
Mosher 1980) but this could simply be due to the non-linear effect of B on P, if
B and age are positively correlated. Others have examined the effect of age on
growth rate. Johnson (1974) found that the growth rate of amphipods declined
with the age of the population, while Coon et al. (1977) found that the
growth rate of mussels increased with age. This contradiction is probably due
in part to the sort of growth rate under discussion. Sutcliffe et al. (1981)
suggest that specific growth rates (% wet wt. day ~') decrease with increased
age, while absolute growth rate (wet wt. day"1) occurs when the animal's
body size is about one-half of the maximum. Although age and biomass are
sometimes confounded, Borkowski (1974) feels that, at least for marine snails,
older populations tend to have higher rates of secondary production. The
lifespan of animals has a similar effect, such that longer-lived animals have
lower rates of production (Zaika 1970; Waters 1977; review by Banse &
Mosher 1980).
The effect of voitinism (number of generations per year) is consistent and
continuous with the effect of lifespan. All authors who cite this effect (e.g.
Johnson 1974: Zytkowicz 1976: Waters 1977: Jonasson 1978; Banse &
Mosher 1980; Benke & Wallace 1980; Wildish & Peer 1981) suggest that
secondary production and P/B increase with the number of generations
Assessment of Secondary Production: The First Step
7
produced per year. Populations that are multivoltine have higher rates of
production than those that are univoltine. An analysis presented by Jonasson
(1978) suggests that we may have erroneously ascribed causation in this
apparent correlation. He suggests that faster growth in the littoral zone
permits more generations per unit time. Thus, multivoltinism may be an effect
of high production rates, not a cause of them.
3.1.3 Body-size
The effect of body-size on secondary production is one of the few relationships
that have been tested explicitly. Unfortunately, much of this work has
employed P/B as a dependent variable and is, therefore, difficult to interpret
mechanistically. The conclusion has been that P/B decreases with increasing
body-size (M) in the population (Janicki & DeCosta 1977; Waters 1977;
Finlay 1978; Banse & Mosher 1980; Benke & Wallace 1980). Banse & Mosher
(1980) have shown that P/B varies as a function of M:
P/B = aMb
(1.3)
where a and b are fitted constants. Because B = NM(N = average population
density) then:
P = aNcM1+b
(1.4)
where c = 1. This equation suggests that the effect of body-size would be more
accurately determined by a multiple regression employing both population
density and mean body-size (see Chapter 8). There appears to be a real effect of
body-size on secondary production, upheld by the experiments of Zelinka
(1977) who found that benthos communities made up of larger species had
lower overall rates of secondary production.
3.1.4 Taxonomy and trophic status
A variety of authors have suggested that physiological and ecological
differences among taxonomic units account for differences in secondary
productivity. Jonasson (1978) suggests that similar species have developed
different tolerances and efficiencies for dealing with environmental problems,
thus production rates must vary among species. Coon et al. (1977) suggest the
same for mussels. Makarewicz & Likens (1979) suggest that differences in P/B
for rotifers among lakes are probably due to taxonomic differences. A number
of workers (Mikulski et al.
1975; Pederson et al.
1976; Waters 1977;
Nauwerck et al. 1980) have suggested that cladocerans are more productive
than copepods, which are, in turn, more productive than rotifers. Schindler
(1972), however, suggests that P/B is higher for rotifers than for other
8
Chapter 1
plankton, thus the apparent low productivity of rotifers could be due to
inaccurate biomass estimation. Herbivorous taxa are generally thought to be
more productive than detritivores or carnivores (Waters 1977; Jonasson
1978).
3.2 Effect of environmental factors
It is one of the basic tenets of ecology that the success of organisms in a
particular ecosystem is determined in
part
by the suitability of the
environment. Among the most obvious aspects of the environment that might
affect animal production are the average temperature, the ability of the
ecosystem to produce sufficient food of acceptable quality, the character of the
substrate, and the concentration of respirable oxygen.
3.2.1 Temperature
Temperature has long been known to influence rates of activity from a
molecular to an organismal scale. It is not surprising, therefore, that many
production ecologists have found that rates of secondary production increase
with temperature (e.g. Neves 1979: Laville 1971; McNaught & Fenlon 1972;
Edmondson 1974; Kititsyna & Pidgaiko 1974; Paterson & Walker 1974;
Pederson et al. 1976; Zytkowicz 1976; Iverson & Jesson 1977; Finlay 1978;
Selin & Hakkari 1982). P, B also is thought to rise with increased temperature,
either as a linear (Winberg et al. 1973; Johnson 1974; Paterson & Walker
1974; Wildish & Peer 1981; Uye 1982) or a curvilinear (Johnson & Brinkhurst
1971;Janicki&DeCosta 1977; Waters 1977; Nauwerck etal. 1980) function.
Banse & Mosher (1980), on the other hand, show that P/B is not correlated
with temperature after regression on body-size.
The general positive effect of temperature on secondary production is a
result of the reproductive biology of zooplankton and benthos. A variety of
authors have suggested that growth rates increase (Johnson 1974; Jonasson
1978: Humpesch 1979; Vijverberg 1980: Marchanl & Hynes 1981; Sutclifle et
al. 1981), egg development times decrease (Schindler 1972: Bottrell 1975;
Makarewicz & Likens 1979: Vijverberg 1980), the rate of population increase
rises (Armitage et al. 1973), and feeding rates increase (Zimmerman & Wissing
1978; see Chapter 9) with increased temperature. These factors tend to
increase production at high temperature (see Chapter 2). On the other hand,
O'Brien et al. (1973) suggest that average clutch size of Diaptomus leptopus
decreases with temperature, and Aston (1973) suggests that egg production by
oligochaetes declines at high temperature. Pidgaiko et al. (1972) conclude that
temperature variation could have either a positive or negative effect on
secondary production, depending upon geographic location and basin
morphometry.
Assessment of Secondary Production: The First Step
9
3.2.2 Food production, availability, and quality
A community of heterotrophs can fix no more energy than the amount made
available to them by primary producers. Edmondson (1974) has reasoned that
the rate of primary production must set the upper limit for secondary
production. Using similar logic, many authors have suggested that rates of
production of freshwater benthos and zooplanklon are positively related to
food availability (Miller et al. 1971; Ladle et al. 1972; George & Edwards
1974; Prikhod'ko 1975; Martien & Benke 1977; Jonasson 1978: Neves 1979;
Benke & Wallace 1980; Nauwerck et al. 1980). Others have found that rates of
zooplankton and benthos production are positively related to rates of primary
production (Patalas 1970, cited by Schindler 1972; Hillbricht-Ilkowska 1972,
cited by Pederson et al. 1976; Monokov & Sorokin 1972; Brylinsky & Mann
1973; Johnson 1974; Dermott et al. 1977; Makarewicz & Likens 1979; Smyly
1979: Brylinsky 1980; Strayer et al. 1981). Winberg (1971b) has been more
specific, hypothesizing that secondary production (Ps) is about 10% of
primary production (Pp), on the average. This suggests that:
Ps = a + bPp
(1.5)
where a = 0 and b = 0-1. A recent analysis by Brylinsky (1980) shows that
phytoplankton primary production is a better predictor of zooplankton
production than phytoplankton biomass, but the relationship may not be
linear. Equation 1.5 probably overestimates zooplankton production at low
phytoplankton production, and makes underestimates at high phytoplankton
production.
The
relationship
between
phytoplankton
production
and
secondary production is probably also responsible for apparent relationships
between secondary production and nutrient conditions (e.g. Stross et al. 1961;
Halle/ al. 1970; Wattiez 1981) and alkalinity (Waters 1977; Pinel-Alloul 1978;
Neves 1979). It should also be remembered that quality of food is important in
determining the secondary production of both zooplankton (Pederson et al.
1976; Vijverberg 1976, 1980; Makarewicz & Likens 1979; Nauwerck et al.
1980), and benthos (Swiss & Johnson 1976; Willoughby & Sutclifle 1976;
Zimmerman & Wissing 1978; SutcliflTe et al. 1981).
3.2.3 Oxygen concentration
The availability of oxygen is thought to be critical, especially to the benthos
because they often live in areas that are oxygen-poor. Brylinsky (1980),
however, has found that carnivorous zooplankton production in a wide range
of lakes is also influenced by oxygen concentration in the epilimnion.
Jonasson (1978) suggests that sufficient oxygen is important to benthos
production because food cannot be metabolized efficiently at low oxygen
levels. This conclusion has also been reached by Dermott et al. (1977) and
10
Chapter I
Rosenberg (1977). Aston (1973) suggests that egg production in freshwater
oligochaetes is constant with decreasing oxygen concentration until some
critical low level is reached. Pond benthos seem to require >lmgl~1 of
dissolved oxygen in order to maintain positive production (Martien & Benke
1977). Laville (1971) suggests that, at least for some benthos, secondary
production and oxygen concentration are inversely related (see also regression
analysis of Brylinsky 1980).
3.2.4 Substrate characteristics
Another aspect of the environment that has been hypothesized as important to
lake and stream benthos is the character and composition of the substrate.
Resh (1977), for example, found that the production of stream caddisflies was
positively related to the average size of particle in the substrate. Hamill et al.
(1979), working in a large river, found that the production of benthic snails
was highest at intermediate substrate particle size. Similar suggestions have
been made by Mecom (1972), Martien & Benke (1977), and Neves (1979). For
lacustrine benthos, secondary production seems to rely more heavily on
organic matter content than particle composition (e.g. Johnson
1974;
Zytkowicz 1976; Marchant & Williams 1977; Jonasson 1978). In addition,
Zytkowicz (1976) feels that benthos production in lakes is a positive function
of the depth to which sediments can be penetrated by benthic organisms.
3.2.5 Miscellaneous environmental factors
Three hypotheses have been advanced which do not fit neatly into broader
categories but which are, nonetheless, interesting. An important factor in
streams and rivers seems to be the current velocity. Zelinka (1977), Hamill et
al. (1979). and Neves (1979) all suggest that secondary production decreases
with increasing water flow rate. With respect to lacustrine zooplankton
production, Edmondson (1974), Makarewicz & Likens (1979), and Selin &
Hakkari (1982) have suggested a positive relationship with intensity of solar
radiation. Finally, Burgis (1971) and Paterson & Walker (1974) suggest that
high zooplankton and benthos production rates should be found in the most
stable ecosystems.
3.3 Predation, competition, and diversity
Predation, competition, and diversity are three topics that have generated
much interest in ecology, yet production biologists have seldom considered
them. Current thought regarding the effect of predation upon secondary
production is contradictory. Hall et al. (1970), Zytkowicz (1976), Waters
Assessment of Secondary Production: The First Step
11
(1977), and Banse & Mosher (1980) suggest that predation leads to increased
production, presumably because the slow growing organisms are removed
from the population. Zndanova & Tseyev (1970), Miller et al. (1971),
Prikhod'ko (1975), and Momot (1978) suggest that predation decreases
production perhaps due to a decline in growing biomass. Thoughts on
competition are less contradictory but less well developed. The basic belief is
that competition decreases the production of a population (see George 1975;
Benke 1976; Lavandier 1981). Production ecologists have not considered the
possible positive effects of competition on community production (cf.
economic theory). The effect of diversity upon secondary production has only
been considered (to my knowledge) by Paterson & Walker (1974). Their data
suggest that the low benthos diversity in a saline lake allowed very high rates of
secondary production.
3.4 Lake morphometry, lateral zonation, and allochthonom input
The morphological characteristics of the ecosystem or placement within it also
seems to affect secondary production. The literature generally suggests that
shallower lakes support higher rates of secondary production (Johnson 1974;
Zytkowicz 1976; Matuszek 1978; Brylinsky 1980). Johnson (1974) also
suggests that the surface area of a lake may be important, since in larger lakes
the profundal zone is less enriched by the littoral zone or allochthonous
sources. Other authors have suggested the importance of allochthonous
materials to secondary production in both lakes and streams (Edmondson
1974; Willoughby & Sutcliffe 1976; Marchant & Williams 1977; Martien &
Benke 1977; Waters 1977; Adcock
production in the littoral zone, it
production in near-shore areas and
other areas (Mathias 1971; Johnson
1979). Possibly due to high primary
is generally believed that secondary
macrophyte beds is greater than in all
1974; Kajak & Dusoge 1975a,b, 1976;
Mikulski et al. 1975; Jonasson 1978; Neveau & Lapchin 1979; Kajak et al.
1980). The only contradiction seems to be for some stream ecosystems where
highest rates of productivity are seen in mid-stream (e.g. Neves 1979).
4 Concluding Comments
The preceding paragraphs indicate that many variables are involved in the rich
variety of hypotheses regarding secondary productivity. In some cases, it is
difficult to extricate real effects from artefacts. For this reason tests of
hypotheses should take one of two courses. Either we should test for the effect
of certain factors under conditions that control all other variables, or we must
pose multivariate hypotheses that account for simultaneous covariation in
more than two variables. I believe that the former approach is currently more
12
Chapter 1
popular because it is conceptually simple; while the latter approach is more
useful, because it is difficult to control circumstances without altering them.
What is really important, though, is that production ecologists define
problems before seeking methods for their examination. To quote again from
Northrop (1947): 'It is like a ship leaving port for a distant destination. A very
slight erroneous deviation in taking one's bearings at the beginning may result
in entirely missing one's mark at the end regafdless of the sturdiness of one's
craft or the excellence of one's subsequent seamanship.' This first chapter has
examined the range of production hypotheses currently under consideration
by production biologists. The chapters that follow strive to supply methods
that can be used to test these and other production hypotheses.
5 References
Adcock J.A. (1979) Energetics of a population of the isopod Asellus aquaticus: life
history and production. Freshw. Bioi, 9, 343-355.
Allen K.R. (1971) Relation between production and biomass. J. Fish. Res. Board Can.,
28, 1573-1581.
Armitage K.B., Saxena B. & Angino E.E. (1973) Population dynamics of pond
zooplankton, I. Diaptomus pallidus Herrick. Hydrobiologia, 42, 295-333.
Aston R.J. (1973) Field and experimental studies on the effects of a power station
effluent on Tubificidae (Oligochaeta, Annelida). Hydrobiologia, 42, 225-242.
Banse K. & Mosher S. (1980) Adult body mass and annual production/biomass
relationships of field populations. Ecol. Monogr., 50, 355-379.
Benke A.C. (1976) Dragonfly production and prey turnover. Ecology, 57, 915-927.
Benke A.C. & Wallace J.B. (1980) Trophic basis of production among net-spinning
caddisflies in a southern Appalachian stream. Ecology, 61, 108-118.
Borkowski T.V. (1974) Growth, mortality and productivity of south Floridian
Littorinidae (Gastropoda: Prosobranchia). Bull. Mar. Sci., 24, 409-438.
Bottrell H.H. (1975) The relationship between temperature and duration of egg
development in some epiphytic Cladocera and Copepoda from the River Thames,
Reading, with a discussion of temperature functions. Oecologia, 18, 63-84.
Brylinsky
M.
(1980) Estimating the
productivity of lakes and reservoirs.
In
E.D.LeCren & R.H.Lowe-McConnell (eds.), The Functioning of Freshwater
Ecosystems. IBP 22. Cambridge: Cambridge University Press.
Brylinsky M. & Mann K.H. (1973) An analysis of factors governing productivity in
lakes and reservoirs. Limnol. Oceanogr., 18, 1-14.
Burgis
M.J.
(1971)
The
ecology
and
production
of copepods,
particularly
Thermocyclops hyalinus, in the tropical Lake George, Uganda. Freshw. Biol., 1,
169-192.
Burke M.V. & Mann K.H. (1974) Productivity and production to biomass ratios of
bivalve and gastropod populations in an eastern Canadian estuary. J. Fish. Res.
Board Can., 31, 167-177.
Clarke G.L. (1946) Dynamics of production in a marine area. Ecol. Monogr., 16,
321-335.
Coon T.G., Eckblad J.W. & Trygstad P.M. (1977) Relative abundance and growth of
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