OXYGEN CONSUMPTION IN D1APTOMUS Gabriel W. Comita Department of Zoology, North Dakota State University, Fargo 58102 ABSTRACT The oxygen consumption of fed and unfed Diaptomus siciloides, D. oregonensis, D. Zeptopus, and D. &wipes was measured at five different tcmpcraturcs using a microWinkler technique. Regression relationships between log oxygen uptake and each of the following independent variables, tcmperaturc, log length, and log weight, as well as the combinations of temperature and length and of temperature and weight, were examined for the four species mentioned and Diaptomus arcticus and Mixodiaptomzrs Zaciniutus. The simple regression of log oxygen consumption on log dry weight at the five tcmpcratures and the multiple regression of log oxygen consumption on temperature and log dry weight gave close approximations to the cxperimcntal means. Kubik Eor her help during the experimental phase of this study. I am especially gratcful to D. T. Eastlund for his extra care and diligcncc during all phases of the study. Parts of this work wcrc supported by the North Dakota Institute for Regional Studies and by National Science Foundation Grant 16415. INTRODUCTION Few oxygen consumption measurcmcnts have been made with the freshwater calanoids of the genus Diaptomus since the work of Scherbakoff ( 1935). For those that have been recorded, the nutritional state was usually mlknown, which probably contributed to the great amount of variability reported. In this study, respiration measurements were made at four or five different temperatures, in the range between 5 and 25C, for five species of female Diaptomus ranging in cephalothorax length from 0.80 to 3.11 mm. The species, all collected near Fargo, North Dakota, were Diaptomus siciloicles Lilljeborg 1889, D. oregonensis Lilljeborg 1889, D. Zeptopus S. A. Forbes 1882, D. clavipes Schacht 1897, and D. arcticus Marsh 1920. In addition, rcspiration measurements: for Mixodiaptomus Zaciniatus Lilljeborg 1889 from Lago Maggiorc, Italy (Comita and Comita 1964), were compared with those for the five local organisms. For several species, some individuals were fed and some were not fed before the experiments to demonstrate how oxygen uptake was affected by feeding. The effect of temperature on the rclationship between oxygen consumption and length or weight was also examined. I am grateful to L. D. Zinsli and J, E. Schindler for their assistance while they were National Science Foundation undergraduate research participants and to P. J, MATFXALS AND METHODS Most of the animals were taken by towing a No. 12 net from short; only D. siciloides was taken in open water. The organisms wcrc sorted into groups of up to five per IO-ml beaker of water from the same source that had been membrane filtered. This membrane-filtered water was used for conditioning and for the measurements of respiration, after aeration at the proper temperature until oxygen saturation reached near 90%. Reforc each experiment, about 40 lo-ml beakers were set up and females of one species were segregated into each beaker, Half the females received no food and the others were given a suspension of Chlamydomonas eugametos sufficient to provide a concentration of 50,000 cells/ml. Nonovigerous females were usually selected for the experiments, but at times it was necessary to include some bearing eggs. Appropriate numbers of these animals were then conditioned at a specified temperature for from 15 to 70 hr ( Table 1). At the end of this period the gut of the fed animals 51 52 GABRIEL W. COMITA 1. Oxygen consumption data for fed and starved females and some males of five species of Diaptomus and one of Mixodiaptomus and the results of Bartlett’s test of homogeneity of variances. Observations were not made at 5C for D. siciloides and M. laciniatus nor at 1OC for D. leptopus and D. clavipcs. D. arcticus and M. laciniatus were not fed during the conditioning period. The variances for D. leptopus are homogeneous at the 5% level TABLE Species D. siciloides D. oregonensis Mean length cephalothorax (mm) ?ep Fed Starved c.v.* (%) (1) (2) (3) (4) (5) (6) (7) (8) D. leptopus D. clavipes D. arcticus Conditioned (hr) Incubated ( hr) 10 15 20 23.5 7” 6 13 5 5 4 3 21 16 19 20 5.049 1.8-2.9 2.5-3.4 3.8-6.0 0.017 0.049 0.075 0.090 0.008 0.043 0.064 0.074 78.5 44.0 67.9 26.4 7 6 5 7 7 5” 4 3 15 19 16 20 18 17 4.9-6.0 5.7-6.7 2.5-3.6 3.0-4.2 5.2-6.2 2.7-3.7 0.023 0.026 0.042 0.041 0.064 0.098 0.023 0.023 0.054 0.028 0.042 0.084 39.5 38.8 50.8 39.6 26.3 31.7 10 15 20 12 10 12 6-7 $ 4$ 3-4 $ 23 23 23 8.4-9.3 4.1-6.5 4.0-4.9 0.021 0.050 0.076 30.8 29.4 20.3 5 15 20 25 9 9 10 10 5 3 2 1 17 17 17 16 5.9-7.1 3.6-4.7 6.2-7.4 4.1-5.3 0.055 0.111 0.178 0.269 0.042 0.079 0.148 0.218 71.5 40.0 23.1 28.6 5 15 20 25 5 15 11 13 3 2 2 1 17 16 18 17 5.7-6.7 3.4-4.9 5.5-6.8 3.5-4.8 0.049 0.150 0.211 0.330 0.031 0.149 0.202 0.330 60.3 21.8 9.5 20.2 5 10 15 20 25 15 14 15 20 15 1 1 1 1 1 21 20 20 70 21 4.3-5.4 5.6-6.7 2.1-3.3 1.3-3.7 1.6-2.7 0.318 0.483 0.634 0.853 1.249 28.9 23.7 26.2 35.5 13.4 H,“,;T;yfvariances X2 df (9) (10) 0.80 1.01 5 10 15 15 20 25 M. laciniatus Females/ vial Oxygen consumed ( pliter 0 -1 hr-1) Determinations 1.02 10.428 1.42 5.87 17.74$ 1.55 25.63$ 3.11 * C.V. = coefficient of variation. t Variances homogeneous between the 5 and 1% levels. $ Variances homogeneous at less than the 1% level. was green, and fecal pellets were present in greater numbers than in the beakers containing unfed animals. After conditioning, the animals were incubated in vials containing approximately 2.2 ml oE filtered and aerated pond water. For the two smaller copepods, D. siciloides and D. oregonmsis, up to six females were enclosed in each vial for the 5 and 1OC measurements; for M. Zaciniatus, six and seven malts were used at 1OC. The number of females per vial was dccreascd according to the size of the organism and as the experimental temperature was increased. For the largest organism, D. nrcticus, only one female was needed to obtain a mcasuremcnt at each of the five temperatures ( Table 1, col. 3). At the end of the incubation period an aliquot was withdrawn from each vial, and the amount of oxygen was determined using the micro-Winkler technique (Barnes 1959). The difference in quantity of oxygen in the experimental vials from that in control OXYGEN CONSUMPTION IN 53 DZAPTOMUS TAULE 2. Regression equations for log oxygen consumption in $ters per female per hr on temperature with ~~~ values computecl from them, and for five species of Diaptomus, and one of Mixodiaptomus, results of tests for deviation from linearity. I?” tests the hypothesis that the regression is linear. Because I?’ was less than the tabular value for I! at the 5% leuel, the hypothesis of linearity for the log expressions is accepted in all cases, All correlation coefficients are significant at the 1% level and all I? ratios are highly significant (OSOJO level) indicating ihat the regression coefficients are different from zero. The ratios listed under F’ are used to test the null hypothesis that the equation is unreliable. F’ is obtained by dividing the regression sums of squares by the residual sums of squares after each is adThe hypothesis is rejected and the equation is reliable if the computed justed for degrees of freedom. F’ is greater than the tabular F (which is not listed in this table). These F’ values are all highly significant indicating reliable equations Test for clcvintion from linearity (logs) D. D. M. D, D. D. n T 32 40 34 38 40 79 0.70 0.76 0.88 0.83 0.89 0.84 Eqnntion Species siciloides log 02 = 0.0574 ( To ) - 2.389 oregonensis log 02 = 0.0342( T”) log 02 = 0.0566( T”) log 02 = 0.0398( T” ) log 02 = O.O431(T”) log Oa = 0.0288( T” ) laciniatus leptopus clauipes arcticus - 1.914 -2.233 - 1.573 - 1.545 - 0.647 vials after the incubation period was the oxygen consumption, expressed as Jiters per female per hr. Length measurements (ccphalothorax) were made on the live females immediately after the oxygen consumption readings, Weight was determined after drying at 1OOC for about 30 min. The data for Mixocliaptomus are from males but appear to bc in the same order of magnitude as that from the females of D. siciloides and D. oregonensis. Bartlett’s test of homogcncity of variances was applied to the data from each species ( Table 1 ), but bccausc the appropriate degrees of freedom in this analysis for all species came to four or less, Box’s modification of this test was also done. The null hypothesis tested by these treatments is that there are no diffcrcnccs in variances in the respiration measurements at each temperature. If the sample variances are very much different from one another, the chi-square and Box’s F-ratio are greater than the tabular values. The results of both tests show that the variances for only one species, D. Zeptopus, arc homogeneous, RESULTS AND DISCUSSION F’ 28.9 53.0 108.7 82.4 146.7 181 QKI 3.75 2.2 3.68 2.5 2.70 1.94 F” 1.52 0.62 3.83 0.06 1.80 0.69 Tabnlnr ( Fo.m = (F0.20 = (F’o.oo = (Fo.,,, = (Fo.lo = (Fo.ro = F df 1.71) 1.64) 4.17) 1.70) 2.49) 1.59) 2,28 3, 30 1, 30 2, 30 2,30 3, 60 uptake were being obtained with D. siciloicles from a lake that contained a large population of Daphnia middendorfiana. At the time of collection the Secchi disc reading was 4.9 m, which was at or near the bottom of the lake. The usual reading during 1965 was 0.3-0.9 m. Because considerably higher respiration rates had been obtained earlier with the same species from the same lake, it seemed possible that the low rates were caused by insufficient food. To test this hypothesis, one group of the females was fed a suspension of C. eugametos containing 50,000 cells/ml while a replicate group was kept without food. Nearly all the readings of oxygen uptake obtained from the fed females were higher than those obtained from the starved ones (Table 1, col. 6 and 7). Similarly designed expcrimcnts were also carried out with D. oregonensis, D. leptopus, and D. clavipes. Respiration rates for D. Zeptopus were also significantly greater for fed than for unfed animals, but in the case of D. oregonensis and D. clavipes the presence or absence of food produced no consistcnt differences, Effect of food Effect of temperature During the course of the measurements it was noted that low readings of oxygen Logarithms of the oxygen consumption rates for the five species of Diaptomus and 54 GABRIEL W. COMITA 3. Log Oa as a function of length at five temperatures. Oxygen consumption is in j.&ters per female per hr and length in mm. All correlation coefficients are significant at the 1% level. All F (= F’ in Table 2) ratios are significant at the 0.5% level TAISLE . A - Temp (Cl Eqlration n 5 log 02 = 0.528L - 2.153 ;/; 10 15 20 25 ~/.A-----+ .u .,.,._................... I.&“---.---l-ec.-- I I 5” I I I I I I S -0 I IO0 I I I I I I 15” I I I I 20” I I I 25” log log log log 02 02 0s 02 = = = = 0.652L 0.521 L0.486L 0.476L- 2.355 1.820 1.594 1.373 30 27 37 43 32 T 0.92 0.95 0.93 0.91 0.79 F 157 228 235 192 346 TEMPERATURE FIG. 1. The oxygen consmnption of five species of Diaptomus over the temperature range from 5 to 25C. The curves are derived from the regression equations presented in Table 2. Closed circles are the means from Table 1, col. G, for oxygen consumption of Diaptomus arcticus, mean cephalothorax length, 3.11 mm (A); open squares, D. &wipes, 1.55 mm (C); small closed circles, D. leptopus, 1.42 mm ( L); small open circles, D. siciloicles, 0.80 mm (S ); and closed triangles, D. oregonensis, 1.01 mm (0). for 121.Zacirz.iatuswere plotted against tcmperature, and curves were fitted to the data by the method oE least squares (Table 2). For all curves, the regression coefficients were significantly different from zero, and there was no deviation from linearity for the semilog plots. Q1o values derived from the equations range from about two for D. arcticus to 3.75 for D. siciloides. Respiration data wcrc also plotted against temperature without transformation to logs. The curves for the five Diaptomus species are shown in Fig. 1. The equations for these curves were also tested Eor nonlinearity. In the case of the three larger species ( curves A, C, and L ), the temperature-oxygen relationship was not linear. Relationship between respiration and size The second regression relationship examined was that of log O2 vs. length for each of the five temperatures; these equations arc listed in Table 3. (The values listed in Table 5, col. 3, arc obtained when the equations are solved at different temperatures.) The F-ratios that were derived from the equations are highly significant and are approximately twice those obtained for the data involving temperature. The calculated means from these equations are closest to the experimental means for M. laciniatus (see Table 5, col. 1). The regression equations of log oxygen consumption on log dry weight for each of the five temperatures as well as for all tem- TABLE 4. Regression equations for log oxygen consumption and log weight in log form and in exponential form. Log Oa is expressed as pliters consumed per mg per hr; log weight as pg dry weight. For each species the mean weights were: D. siciloides, 3.2; D. oregonensis, 4.75; D. leptopus, 22.2; D. clavipes, 28.7; D. arcticus, 300. In exponential form, the R is oxygen consumption in ,uliters per female per hr. All correlation coefficients are significant at the 1% level. All l? (= F’ in Table 2) ratios are significant at the 0.5% level Tknp Equation (C) 5 10 15 20 25 All 1: Standard log log log log log log error 02 = Oa = 02 = O:! = 02 = 02 = 0.81310.8618 1.1714 1.3938 1.5775 1.1635 of cstimatc - n 0.3314 0.2792 0.3462 0.3743 0.3780 0.3418 of y (log 0,). (log (log (log (log (log (log W) W) W) W) W) W) 5 5 5 5 25 T -0.97 -0.98 -0.98 -0.99 -0.99 -0.67 SE* 0.065 0.055 0.020 0.019 0.028 0.272 F 63 64 698 899 441 19 Exponential R = R = R = R = R = R = form 6.50w”~ooD 7.27 W”.721 14.84 W”.O”l 24 76 W”.Ozo 3i.80 W”.“‘” 14.57W”J”” OXYGEN TABLE 5. peratures. Te111p (C) CONSUMPTION IN Comparison of the results obtained from the five different regression equations at five temP are the experimental means for oxygen consumption as given in Table 1, col. 6. Entries in all columns are in pliters of oxygen per female per hr P Temp Length (1) (2) (3) (2) (2) and (3) nnd (6) (4) (5) 0.019 0.015 0.040 0.062 10 15 20 25 0.011 0.021 0.022 0.050 0.041 0.076 0,079 0.152 0.024 0.020 0.051 0.080 0.130 5 10 15 20 25 0.049 0.150 0.211 0.330 0.045 0.045 0.097 0.144 0.232 10 15 20 23.5 25 - 0.027 0.038 0.053 0.073 - Tcmp Length and (3) nnd (6) Log W (6) (1) (2) (3) (4) (5) (6) D. oregonensis 0.013 0.021 0.031 0.048 0.014 0.017 0.032 0.051 0.102 0.103 0.073 0.078 0.034 0.048 0.06G 0.092 0.128 0.023 0.026 0.042 0.064 0.018 0.027 0.040 0.059 0.062 0.087 0.120 0.165 0.235 0.024 0.020 0.051 0.079 0.033 0.047 0.064 0.090 0.018 0.027 0.041 0.062 0.018 0.022 0.041 0.066 0.098 0.087 0.128 0.122 0.094 0.099 D. leptopus - - 0.055 0.111 0.178 0.269 0.047 0.075 0.119 0.187 0.296 0.040 0.037 0.083 0.125 0.201 0.058 0.087 0.133 0.202 0.307 0.061 0.082 0.133 0.202 0.305 0.318 0.483 0.634 0.853 1.249 0.313 0.437 0.609 0.848 1.181 0.308 0.471 0.631 0.827 1.281 0.054 0.075 0.105 0.145 0.205 0.048 0.073 0.111 0.169 0.257 0.051 0.068 0.112 0.171 0.259 0.269 0.409 0.622 0.945 1.436 0.366 0.444 0.618 0.878 1.313 D. arcticus D. clauipes 0.047 0.077 0.126 0.208 0.341 (2) 9 M. laciniatus 5 (2) Log w D. siciloides 0.008 0.017 0.015 0.049 0.030 0.075 0.057 0.090 (0.10) 0.111 5 55 DIAPTOMUS pcratures combined arc prescntcd in Table 4 and shown graphically in Fig. 2. Solutions based on these equations are listed in Table 5, col. 6. These solutions are the closest overall approximations to the cxpcrimcntal means obtained for the six spccics, although the temperature equations (Table 2) seem to bc equally acceptable. The slope of the 1OC solid lint (Fig. 2) deviates most, but not significantly (F = 1.99, df 4, 15; Fo.05= 3.06) from that of the lines at the other temperatures. If these equations arc in fact a family of curves as they appear to be, this deviation may bc enough to yield low rates of oxygen uptake in the lower weight lcvcls shown in Fig. 2. When these equations are transformed from the log form to the exponential form, the constants obtained range from 6.622 to 0.721. The exponential constant (0.658) from the equation combining data from all temperatures can bc compared with those given by Conover (1960) : for herbivores, 0.559, and for carnivores, 0.649; although Conovcr’s data were obtained over a much 0.339 0.480 0.640 0.909 1.264 narrower range of temperature. I?or the five species of Diaptomus, this constant is closer to the theoretical value, expressed by the relationship R = hW0.67, relating respiration to surface area or % power of weight. There is no evidence that any of thcsc diaptomids are carnivorous. The data presented by Raymont and Gauld ( 1951)) Gauld and Raymont ( 1953)) Conover (1956, 1959,196O) and Raymont ( 1959 ) for scvcral marinc copepods can be compared with these for Diaptomus. Conovcr’s analysis led him to accept a direct rcla tionship, involving no transformations, bctwecn oxygen consumption as pliter per 1,000 copepods per hr and mg dry wt per 1,000 copepods. The equivalent equation for these Diaptomus species based on weight is 02 = 2.1466 (wt as mg) + 0.06744. Since size measurements involve either length or weight or both, examination was made of the lengths and weights available for five of the six species studied and additional length-weight data were used that were taken from scvcral other local 56 GABRIEL W. COMITA TARLE 6. Ponderal index values, k, derived from length (mm) and weight (pg) data taken from 500 or more specimens of the seven species listed Length (mm) Species D. siciloicles D. oregonensis D. sicilis II). saskatchewanensis leptopus clavipes arcticus arcticus (1965) D. D. D. D. FIG. 2. Rcgrcssion lines for oxygen consumption as a function of dry weight at five tempcratures. Lines arc dcrivcd from the equations in Table 4. Data points plotted arc the experimental means shown in Table 1, col. 6, except for 5C at 3.2 ,ug and 1OC at 22.2 and 28.7 ,ug. The 5C point is the cxpcrimental mean ( closed circles ), 4.7 pg mg-’ hr-I, from unfed D. siciloides. It is the mean respiratory rate of nine replications with seven females in each vial. The 1OC points (closed triangles) plotted at 22.2 ,LL~ (D. Zeptopus) and 28.7 ,ug (D. &Apes) as well as those for 3.2 ,ug (D. siciloides) at 15 and 20C were derived from equations in Table 2. The 15C line (dashed) represents two nearly identical lines, one for the 15C data and one for all temperatures combined; they are not separable in the figure. populations, including some from these same species taken in 1963, (Comita and Schindler 1963). The expression, wt (pg) = k ( length3), for the relationship between length and weight as derived from the data on hand was cxamincd (Table 6). The only 7cvalue that departs markedly from the others for the same sex is that for D. nrcticus females ( 1965), which was based on 79 length measurements but on only 15 weight measurements. These femaics had a longer mean cephalothoracic measurement (3.11 mm) than those taken from the sami pond in an earlier year ( 1962). The reason for this difference is not known. The other two 7cvalues for D. arcticus prcsentcd in the table, one for males ( 9.05) and one for females ( 5.84)) are from the 1962 populations based on 500 - specimens, and they are probably more representative because of the larger number weighed. 7c values calculated for Limno- d 0 0.75 0.74 0.80 0.76 1.45 1.39 2.34 0.95 0.78 0.95 0.87 1.70 1.68 2.80 3.11 k d” 9.22 8.32 8.02 10.18 10.04 10.48 9.05 0 5.72 4.94 5.38 6.47 5.52 6.56 5.84 9.73 calanus johanseni (Comita 1956) are about the same as those listed in Table 6. Relationship between respiration and the combination of temperature and size When both length and temperature, including 182 sets of data, are taken as independent variables, a multiple regression equation can be computed to show their combined effect on log oxygen for all the species. The equation, which also yields close approximations to the measured respiration values, is: log O2 = 0.4732 ( L ) + 0.0286 (To) -2.085; (r = 0.80 for log 02 on length; r = 0.35 for log O2 on temperature ). The F-ratio is 116, and again is highly significant. The standard error of Y is reduced (from 0.315 to 0.246) by about 22% by the addition of temperature as the second independent variable. The values obtained from the solution of this equation are shown in Table 5, col. 4. Bimodality of length in Calanoid copepods is a common phenomenon. In these diaptomids, a pronounced bimodality occurred in both the D. siciloides and D. uregonensis populations, and it is certain to be an important factor Mlucncing the precision of an cxprcssion involving length mcasuremerits. For D. siciloides, the cephalothoracic measurements ranged from 0.73 mm to 1.06 mm with the lower mode centering around 0.78 mm and the upper mode centcring around 0.98 mm. For D. oregonensis, a lower mode centers around 0.94 mm and the upper mode around 1.06 mm (absolute OXYGEN CONSUMPTION range, 0.80 to 1.17 mm). For D. leptopus, there was a single mode at I.41 and for D. clauipes and D. arcticus length measurcments were unimodal also. Some of the specimens were weighed at the time of these mcasurcments, and weight recorded at other times from populations of the same spccics from the same ponds were used to supplement the data. The multiple regression equations, combining temperature and all the weight data, were: log O2 = 0.0364 (To) - 1.818 (W) + 0.3068; r = 0.73 for log O2 on temperature; -0.59 for log 02 on weight. IN 57 DIAPTOMUS If one adult female D. middendorfiana. siciloides having 0.028 calories respires at its 20C starved rate, which is 0.064 ,Jiter of Oz/hr (86% of its fed rate), it could theorctically consume itself in respiration in about 3.7 days (assuming no food intake), but could survive for 8.1 days if its respiration dropped to 40% of its fed rate. Thus, a change in respiratory rate from 86 to 40% of the fed rate increases the survival time by a factor of about 2.2. This additional time could help the animal survive a period of intense competition with any herbivore. r= REFERENCES H. 1959: Apparatus and methods of oceanography. Intcrscience, New York. 341 p. CCIMITA, G. W. 1956. A study of a Calanoid lake. copepod population in an Arctic Ecology, 37 : 576-591. -, 1965. The oxygen consumption of Diaptomus sicizooicles, Lilljeborg. Limnol. Oceanog., 10 : 466-468. AND J. J. COMITA. 1964. Oxygen upta<e in Mixodiaptomus laciniatus Lill. Mem. 1st. Ital. Idrobiol., 17: 151-166. -, AND D. W. SCIIINIILE~. 1963. Calorific values of microcrustacea. Science, 140 : 1394-1396. CONOVER, R. J. 19S6. Oceanography of Long Island Sound 1952-1954. VI. Biology of Acartia clausi and A. tonsa. Bull. Bjnghnm Oceanog. Collection, 56 : 156-233. -. 1959. Regional and seasonal variation in the respiratory rate of marinc copepods. Limnol. Oceanog., 4: 259-268. -. 1960. The feeding behavior and respiration of some marine planktonic Crustacca. Biol. Bull., 119: 399-415. GAULD, D. T., AND J. E. G. RAYMONT. 1953. The respiration of some planktonic copepods. II. The effect of temperature. J. Marine Biol. Assoc. U.K., 31: 447-460. RAYMONT, J. E. G. 1959: The respiration of some planktonic copepods. III, The oxygen requirements of some American species. Limnol. Oceanog., 4: 479-491. -, AND D. T. GAULD. 1951. The respiration of some planktonic copepods. J. Marine Biol. Assoc. U.K., 29: 681-693. RICHMAN, S. 1958. The transformation of energy by Daphnia pulex. Ecol. Monographs, 28: BAJXNES, log O2 = 0.0364 (To)-0.3418 + 0.6182; (log W) r = 0.73 for log O2 on temperature; r = -0.67 for log O2 on log weight. In both equations, log 02 is measured in ,Jiter mg-l hr-I, W in the first equation is in mg and in the second in pg, and To is in C. These equations are derived from 25 sets of observations. The F-ratio for the first cxpression is 61 (significant at the 0.5% level) and that for the second is 633-also highly significant. The standard error of the depcndent variable is reduced from 0.253 to 0.050, and the improvement F obtained by the addition of log W as pugis 586. Evaluations of only the equation using the log transformation of weight arc listed (Table 5, col. 5). The log W equations ( Table 4) were solved using the weight and respiration data of other copepods compiled and reported by Richman ( 1958, his Table 7). The oxygen consumption values thus obtained are about 1.2-1.4 times greater than the observed values recorded. Although the nutritional condition of those listed was not known, it is probable that the reported values are low because they are rates from unfed animals. It was mentioned earlier that rates for D. siciloides were consistently lower when living under sevcrc competition with D. s,,E2z-yF: A P 1g35 Ueber den Squerstof. . . L fverbraudh von einigen Planktoncrustacccn. Arb. Limnol. Sta. Kossino, 19: 67-89.
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