production in early summer; in the central and northern portions of 

 their areas of distribution the maximum production is shifted to the 

 heat of summer, approaching the mean annual values in early fall. In 

 Arctic waters, boreal-arctic species yield the maximum production at the 

 end of hydrologic summer. Some high Arctic and Antarctic species show 

 their maximum production in winter. 



In water areas differing in their physical and chemical modes, the 

 production process in populations of the same species occurs 

 differently. Its rate increases with an increase in temperature (within 

 the limits of the optimum for each species), while the degree of 

 fluctuation increases with an increase in the variability of natural 

 conditions. Thus, in the inlets of Pos'yeta Bay, a number of relatively 

 warm-water low-boreal species reach their maximum production in 

 midsummer, while in open areas of the bay, the maximum is reached only 

 in the fall (Golikov, Menshutkin, 1973). Species which are subtropical 

 in origin achieve high production in the boreal waters of the Pacific 

 only in shallow inlets which are thoroughly heated in the summer (even 

 if they are quite remote from each other), and are encountered only as 

 individuals or are completely absent in nearby open, colder sections. 



It is clear that in order to understand the biologic processes 

 occurring in marine ecosystems, we must do more than study the 

 production capabilities of populations of leading species, or even of 

 entire biocenoses. In order to study the regularities of the cycle of 

 biologic energy in ecosystems of various types, we must know, as a 

 minimum, the consumption of food (C) and its assimilation (1/U) for the 

 various trophic levels, the cost of metabolism (R), the effectiveness of 

 utilization of consumed and assimilated food for growth (K^, K2) and the 

 flow of energy through the population, or the assimilated energy (A = 

 P + R). In the literature, a tremendous quantity of information has 

 been accumulated on these parameters for marine benthic organisms. Data 

 are also available on the calorie content of various groups of marine 

 invertebrates and algae in sufficient quantities for measurement of the 

 biologic processes in identical bioenergetic units — calories. Most of 

 the coefficients necessary for calculation vary within relatively narrow 

 ranges. Thus, for most of the organisms studied, the assimilation of 

 food is usually 60-80% of the diet, the regression factor in the 

 equation of metabolism as a function of weight (R = aW ) averages 0.75- 

 0.80 (Hemmingsen, 1952; Vinberg, 1966). The coefficient of 

 effectiveness of utilization of assimilated food for growth is 

 approximately the same for species with similar dimension-weight and age 

 structures in a given landscape-geographic zone, averaging 0.3-0.4 for 

 populations of macrobenthos in temperate waters. In species in the 

 higher latitudes, this coefficient increases significantly due to the 

 greater quantity of lipids in the food, with their high energy capacity 

 (Golikov, 1975a, b) . Of particular interest for an understanding of the 

 trophodynamic relationships in benthic biocenoses is the study of the 

 ecologic effectiveness of populations. This characteristic is 

 calculated as the ratio of production of a population at the ith trophic 

 level to the production of populations of the i-l-th level, consumed by 

 them and, as Lindeman has shown, this characteristic averages about 

 10%. Preliminary calculations of the transformation of energy in the 



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