In defence of average cohorts

SUMMARY. Hynes & Coleman (1968) proposed a method for estimating benthic secondary production for use with populations in which cohorts cannot be distinguished and for use with unidentified benthos. Hamilton (1969) corrected and refined the method, emphasizing the concept of the average cohort. Zwick (1975) recently suggested the method should not be used since: (1) too many conditions need to be filled for use with unidentified material, and (2) it is strongly dependent on growth patterns. This paper shows that Zwick misinterpreted the concept of the average cohort, and his apparent invalidation of the method due to dependence on growth patterns is erroneous.     

Life cycles, microdistribution and production dynamics of six species of net-spinning caddisflies in a large southeastern (U.S.A.) river

Hydropsyche incommoda. H. rossi, Cheumalopsyche pasella, Mocronema Carolina. Chimarra moslyi , and Neureclipsis crepuscularis were studied on woody substrates (snags) in the lower Savannah River. Distribution of most species biomass was positively correlated with attached macrophytes on snags at medium (25–50 cm s^-1) current velocities. All species were bivoltine at the study site with the exception of M. Carolina which was univoltine. Annual and cohort production estimates (Hynes and removal-summation methods) were obtained for each species at three velocity ranges. Total annual caddisfly production calculated by the Hynes method was 12.6, 41.4, and 22.7 g (ash free dry wt m^-2 of snag surface area at low, medium and high velocities, respectively; 9.6, 34.4, and 19.9 g with removal-summation; and, 11.5, 36.1 and 21.1 g when the Hynes method was applied to discrete cohorts. Higher cohort production was attained by the overwintering generation because of higher terminal instar body weights. Dissolved organic carbon comprised most of the total organic load transported by the river; however, abundant paniculate organic food was also apparently available for filler feeders. Organic and inorganic seston concentrations were highest in winter months and lowest in autumn. Over 90 percent of the seston (by weight) was <105 μm. It is suggested that the variation in catchnet mesh size, coupled with spatial heterogeneity of the snag habitat, allows the caddisflies to partition space, which is apparently the limited resource.     

Habitat and substrate influences on population and production dynamics of a stream caddisfly, Ceraclea ancylus (Leptoceridae)

SUMMARY. Population and production dynamics of Ceraclea ancylus (Vorhies), a leptocerid caddisfly with a univoltine life cycle, five larval instars, and a single cohort population, were examined in Brashears Creek, Kentucky. A systematic sampling approach that included analysis in riffle, pool, and waterwillow bed habitats was used and fourteen sampling transects were taken across Brashears Creek between May 1971 and July 1972. For each of the 70 m² samples taken along a transect, the depth, habitat, and substrate type were recorded. The number of C. ancylus cases and numbers and dry weights of each larval instar were determined. The majority of the C. ancylus population was in the second larval instar in July, the third larval instar in August, the fourth larval instar in September, and the fifth larval instar from October through May. The spring population was 30% of the early instar maximum estimated the previous August. The spatial distribution of C. ancylus reflects a logarithmic pattern with early instar larvae more highly clumped than the late larval instar and pupal populations. Initial standing stock estimates were less than 1% of the standing stock biomass. A production estimate of 9.9 mg/m²·year (dry weight) was determined using the instantaneous growth method, Allen's Curve method, and the Hynes method of estimating secondary production. Turnover ratios (7.0 annual TR, 5.7 cohort TR) were higher than estimates reported as typical for benthic macroinvertebrates because of the low initial to final mean individual weight of the C. ancylus larvae (0.4%). Production estimates calculated for C. ancylus populations in waterwillow bed (4.4 mg/m²·year) and pool (13.7 mg/m²·year) habitats were less than calculated for the riffle population (19.9 mg/m²·year). The annual turnover ratio (10.2) was higher in the pool population than in other habitats because of spring larval migration into the pool from other habitats. A trend of higher production estimates with larger sized substrate particles and increased numbers of particles may reflect the increase in available food sources or attachment sites. The accuracy of production estimates and their applicability to water resource management may be limited by a failure to consider the spatial distributions of the population being sampled and the tendency for sampling procedures to be confined to single habitats.     

Nutrient dependence of phytoplankton production in brown-water lakes with special reference to Lake Päijänne

A method for predicting the mean seasonal chlorophyll a concentration, the mean seasonal in vitro phytoplankton primary productivity per unit volume, the maximum daily production per unit volume and the seasonal integral production in brown-water lakes is presented. The production values can be calculated when the mean annual concentration of total phosphorus and the mean annual colour of the water are known. This method has been developed especially for practical water pollution studies to permit rapid and inexpensive estimates of major biological consequences of changes in effluent loads. The method can be applied for brown-water lakes where phosphorus is the limiting nutrient for primary production.     

The effect of organic solvents on the equilibrium position of enzymatic acylglycerol synthesis

The effect of organic solvents on the equilibrium position of lipase-catalyzed esterification of glycerol and decanoic acid has been investigated. The reaction is carried out in an aqueous-organic two-phase system. In polar solvents, high mole fractions of monoacylglycerol and low mole fractions of triacylglycerol and measured, while in nonpolar solvents, the measured differences in the mole fractions of monodi-, and triacylglycerols are less. There is a good correlation between the ester mole fractions at equilibrium and the log P of the solvent (partition coefficient in n-octanolwater), however, only if the group of tertiary alcohols is excluded. In the plot of the easter mole fractions as a function of the logarithm of hte solubility of water in the organic solvent, the tertiary alcohols can be included; however, in this case other deviations appear.For the prediction of the effect of organic solvents on the ester mole fractions at reaction equilibrium in nondilute reaction systems with a water activity below 1, the program TREP (Two-phase Reaction Equilibrium Prediction) is developed, which is based on the UNIFAC group contribution method. With this model the equilibrium data are essentially predicted from basic thermodynamic data. The required equilibrium constants are estimated from experiments without an organic solvent in the reaction medium. The mole fractions calculated by TREP show the same trends as the experimentally measured mole fractions; however, some variation is observed in the absolute values. These deviations may be due to inaccuracies in the UNIFAC group contribution method. TREP is found to be a correct method to predict within some limits the ester mole fractions at equilibrium for all mixtures of solvents, substrates, and products. The production of monoester can be enhanced in reaction system with a sufficient high concentration of a polar solvent. In experiments with a triglymeto-decanoic acid ratio of 5, almost no di-and triesters can be detected at equilibrium. (c) 1993 John Wiley & Sons, Inc.     

Solvent effects on lipase-catalyzed esterification of glycerol and fatty acids

The lipase-catalyzed acylglycerol synthesis with fatty acids of different chain length is studied. Measured ester mole fractions at equilibrium are compared with calculated mole fractions. For these calculations the computer program TREP (Two-phase Reaction Equilibrium Prediction) is used. This program is based on the UNIFAC group contribution method and is developed for nondilute two-phase reaction systems.With one set of equilibrium constants, namely 1.3, 0.8, and 0.6 for monoester, diester, and triester synthesis, respectively, the equilibrium position of the reaction between glycerol and all saturated fatty acids with a chain length from 6 to 18 and oleic acid (cis-9-octadecenoic acid) can be calculated. Deviations, expressed as the ratio between calculated and measured ester mole fractions, usually were between 0.7 and 1.2. In the presence of solvents, the deviations of the monoester mole fractions were higher and rose up to 3. Without addition of a solvent, the ester mole fractions at equilibrium are dependent on the fatty acid chain length. With the short-chain hexanoic acid, the monoester mole fraction is the highest ester mole fraction, while for the long-chain oleic acid, the diester mole fraction is the highest one. The ester mole fractions become independent on the chain length of the fatty acid with a solvent added in a sufficient high concentration. Both reactions, with saturated and unsaturated C(18) fatty acids, lead to the same equilibrium position. The program TREP is found to make good predictions of the equilibrium amounts of ester and fatty acid. However, systematic deviations arise between measured and calculated amounts of water and glycerol in the organic phase. The calculated water and glycerol amounts are always lower than the measured ones. These deviations seem to be highest in nonpolar media and are probably due to deficiencies in the UNIFAC calculation method. Some preliminary experiments show the effect of the choice of solvent on the reaction rates. In polar solvents, the monoester production rate is enhances by a factor of 1.5 as compared to the reaction rate in a system without solvent. (c) 1993 John Wiley & Sons, Inc.     

The effect of water level fluctuations on a littoral fauna

The littoral fauna of Llyn Tegid, North Wales, in 1968/9 is compared with the results of similar faunal investigations made by Rothwell(née Dunn) in 1951/2 (Dunn, 1953) and Hynes in 1957 and 1959 (Hynes, 1961). The results show the long term effects upon the littoral fauna of the change in mean water level and the increased water level fluctuations produced for a time by the controlled outflow scheme, which came into operation in 1955. In 1961 Hynes recorded big reductions in numbers and variety of invertebrates living in the littoral zone. The reduced water level fluctuations in recent years has resulted in a re-establishment of some of the fauna.     

Community production and respiration by ciliated protozoa in the benthos of a small eutrophic loch

SUMMARY. The results of laboratory experiments on ciliate reproductive and respiratory rates have been combined with field data for ciliate numbers, to produce estimates of production and respiratory energy losses by communities of benthic ciliates. Annual production decreased with increasing depth, and it was estimated to be 345, 149 and 40 J cm⁻² year⁻¹ at three sites in Airthrey Loch, Scotland. A similar trend with increasing depth was recorded for annual respiratory energy losses, being 16, 11 and 4 J cm⁻² year⁻¹ at the three sites. The corresponding annual net production efficiencies (%) were 95.2, 93.1 and 91.5. When net production efficiencies were calculated for each period (the time between successive sampling occasions) as 100 (daily production/daily assimilation), the range extended from 63 to 98. The energetic efficiencies of the ciliate populations are discussed with reference to data in the literature for higher organisms.     

Census vs. effective population size in chinook salmon: large- and small-scale environmental perturbation effects

Population viability has often been assessed by census of reproducing adults. Recently this method has been called into question and estimation of the effective population size (Ne) proposed as a complementary method to determine population health. We examined genetic diversity in five populations of chinook salmon (Oncorhynchus tshawytscha) from the upper Fraser River watershed (British Columbia, Canada) at 11 microsatellite loci over 20 years using DNA extracted from archived scale samples. We tested for changes in genetic diversity, calculated the ratio of the number of alleles to the range in allele size to give the statistic M, calculated Ne from the temporal change in allele frequency, used the maximum likelihood method to calculate effective population size (NeM), calculated the harmonic mean of population size, and compared these statistics to annual census estimates. Over the last two decades population size has increased in all five populations of chinook examined; however, Ne calculated for each population was low (81-691) and decreasing over the time interval measured. Values of NeM were low, but substantially higher than Ne calculated using the temporal method. The calculated values for M were generally low (M < 0.70), indicating recent population reductions for all five populations. Large-scale historic barriers to migration and development activities do not appear to account for the low values of Ne; however, available spawning area is positively correlated with Ne. Both Ne and M estimates indicate that these populations are potentially susceptible to inbreeding effects and may lack the ability to respond adaptively to stochastic events. Our findings question the practice of relying exclusively on census estimates for interpreting population health and show the importance of determining genetic diversity within populations.     

The ecology and production of copepods, particularly Thermocyclops hyalinus, in the tropical Lake George, Uganda

(1) Total copepod numbers per litre in Lake George, Uganda, vary slightly during the course of a year. Numbers increase during the dry seasons June-August and January-February, and decrease during the rains. This variation is more marked in the mid-lake area, where copepod numbers are higher, but the change in numbers is not more than two-fold. By comparison with populations in temperate regions the population in Lake George is stable in size and shows little seasonal variation. (2) The carnivorous cyclopoid Mesocyclops leuckarti forms only 4–21 %, by num-bers, of the total copepod population and does not occur in numbers large enough to be analysed profitably. (3) The bulk of the population is the much smaller Thermoeyclops hyalinus and analysis of this population into the main development stages of its life history shows an almost constant age structure over the lake as a whole, throughout the year. (4) The development time of the eggs, as determined previously (Burgis, 1970), is 1–5 days. This, plus mean development-times of 6 and 11 days for nauplii and copepodites respectively, has been used to determine the daily rate of recruitment from each stage of the life history to the next. The recruitment from nauplii to copepodites and from copepodites to adults are very similar and almost constant. Recruitment from eggs to nauplii is. however, mueh greater, and it thus appears that the major loss to the population is at this eariy stage in the life history. This may well be due to predation by the larvae of Chaohorus. (5) Using this recruitment rate from eggs to nauplii, the potential turnover time of population numbers was calculated to have a mean value of 4–2 days for the lake as a whole. This is similar to maximum summer turnover times found by authors such as Hall (1964) for Daphnia populations in temperate waters. (6) The biomass of individuals at various stages in their life history was deter-mined as carbon, and application of these determinations to the numerical counts was used to calculate the standing-crop biomass in the lake. With a constant population of stable age-structure it is possible to estimate the daily biomass production from the turnover time of numbers. For the population of Thermocyelops hyalinus in Lake George this gives a mean figure of 211 μg C/I/day during 1969–70. (7) Instantaneous birth rates and instantaneous rates of change in population numbers (b and r) have also been calculated for the population of Thermocyclops. The values of r show some variation at one centre site, fluctuating around zero, but when calculated for lake mean figures r is hardly different from zero throughout the year. The values of b also show fluctuations when calculated for one site but when calculated for lake mean figures show no more than two-fold variations, being higher at the beginning of the two rainy seasons. The values of b are not as high as those attained during the summer in some temperate Daplmia populations such as those described by Wright (1965). The values of ZJ tend to be higher inshore than in mid-lake areas of the lake. (8) The instantaneous mortality rate r/has been calculated from b-r. With such very small values of r, d is always very similar to b. Most, if not all, recruitment to the population is counterbalanced by loss, probably predation. (9) At one inshore site the population was analysed and counted in detailed cate-gories, over a period of 10 days. This allowed a more accurate estimation of the standing-crop biomass and thus of production. A mean value of 103 μg C/l/day was obtained for these samples. (10) The mean daily level of production, c. 211 μg C/l/day, o(Thermocyclops hyalinus in Lake George, is among the higher values of zooplankton production available for comparison most of which are from temperate regions of the world. The fact that it is applicable throughout the year gives an annual value of 77g C/m³/year which is probably high.