Author Institution: Department of Physiology, College of Medicine, The Ohio State University, Columbus 1

Angerer, Clifford A.

Murray, Mary C.

The Ohio Journal of Science

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OXYGEN CONSUMPTION OF ADRENALECTOMIZEDAND RENAL DAMAGED FROGS12CLIFFORD A. ANGERER AND MARY C. MURRAYDepartment of Physiology, College of Medicine, The Ohio State University, Columbus 10The frog, both as a representative of poikilotherms and as experimental ma-terial for adrenocortical investigations, possesses certain advantages overhomoiotherms for studies involving penetration rates of known molecular-volumevital dyes and of radioactive physiological ions through its integument (Angerer,1942), of water into and from the whole organism (Angerer and Angerer, 1949;Angerer, 1950) and into and from skeletal muscles (Angerer and Angerer, 1950),and for studies treating of skin bio-electric potentials (Williams and Angerer,1950). Recently success has been attained with injections of adrenocorticalextract (ACE) into otherwise normal frogs with regard to metabolism (Calhoonand Angerer, 1951) and skin bio-electric potentials (Williams and Angerer, 1952).In the light of these studies and of the paucity of data treating of the metabolismof normal frogs, it was considered desirable (a) to determine for the first timethe rates of oxygen consumption at various intervals of time after renal damageand (b) after adrenalectomy, with the view of extending the study of adrenocorticalfunction to the poikilotherms, and (c) to obtain data for the oxygen consumptionof another species of frogs by a different respirometric technique.METHODUnfed male frogs, Rana pipiens, of winter (late autumn and winter) and earlyspring (before spawning) stock, weighing between 20 and 30 gm. and showingtesticular fat at autopsy, were divided into 3 major groups (12-26 frogs/group):(1) adrenalectomized, both "paired" (curve A1, fig. 1) and "unpaired" (curveA), (2) renal damaged (sham-operated control, curve R), and (3) unoperated(normal control, curve N) frogs. The method of adrenalectomy, of renal damage,of postoperative care on sphagnum moss, and of determining adrenocorticalinsufficiency has been described elsewhere (Angerer and Angerer, 1949; Angerer,1950). "Paired" animals (included in groups 1 and 2) served as their own con-trols prior to adrenalectomy; "unpaired" frogs had their controls (unoperatedand renal damage) randomly and simultaneously selected from the same stockas those designated for a given operative procedure.Oxygen consumption was determined by direct-reading volumeters. Theaccurac}^ of this technique is comparable to that of the Winkler method (Calhoonand Angerer, 1950, 1953). The frogs were gradually acclimated (17-21 hr.) tothe temperature of the waterbath to prevent possible thermal stimulation (Angerer,1940). The frogs were permitted to sit in a given volume of tap water sufficientto moisten their integuments. The volumeters were oscillated through a distanceof 3 cm., at a frequency of 110 cycles/min. and at a bath temperature of 30±0.01° C. After a 15-minute equilibration period, readings were made immediatelyat 15-minute intervals for at least one hour. All readings were corrected tostandard pressure and unit body weight. The results were expressed as ml.oxygen consumed/gm. of body weight (wet)/hr.1 Aided by grants from the Comly-Coleman Research Fund, College of Medicine, and fromthe Graduate School, The Ohio State University.2Presented at the Columbus meetings, American Physiological Society: Amer. Jour.Physiol. 163: 737. 1950.THE OHIO JOURNAL OF SCIENCE 55(5): 296, September, 1955.No. 5 OXYGEN CONSUMPTION OF FROGS 297RESULTS AND DISCUSSIONNormal frogs. The mean oxygen consumption for all (38) frogs studied is0.175 ±0.003 (standard error) ml./gm./hr. at 30° C. There is little fluctuationin this mean over 5 successive days as is indicated by the low value for its standarderror. Curve N, figure 1, serves not only for purpose of reference when the percentDAYSPOST-OPERATIVEFIGURE 1. The percent variation in mean oxygen uptake (ml./gm./hr.) for renal damaged(R) and for "paired" (A1) and "unpaired" (A) adrenalectomized frogs compared with normal(N) frogs (ordinates) is plotted against their postoperative time in days (abscissae). "Basal"winter and spring frogs were studied at 30° C. ( "Paired" indicates each frog served as itsown control.)variation in mean oxygen uptake (ordinate) is plotted against postoperative timein days (abscissa) but also indicates, incidentally, the mean oxygen uptake (0.175ml./gm./hr.) of all normal frogs during a period of 5 successive days. Whenwinter and spring frogs are studied, under the stated experimental conditions,298 CLIFFORD A. ANGERER AND MARY C. MURRAY Vol. LVthe means are 0.183±0.006 and 0.165±0.006 ml. 02/gm./hr., respectively. Thedifference (-9.8%) between these values is not significant (P>0.05, Student'smethod); therefore, the mean for all stocks is used in subsequent comparisons.The lack of significance between the means for our winter and spring stocks isprobably due to all frogs being kept under routine laboratory conditions, as pre-viously outlined, and in a "basal" state, in that the frogs were unfed and sub-sisted on their fat depots. The values hitherto reported (Loewy, 1926) for 2frog species, studied at 20° C, are 0.080 and 0.260 ml. 02/gm./hr. for winter andApril R. temporia and 0.100 and 0.210 ml. 02/gm./hr. for winter and April R.fusca, respectively. Our mean of 0.175 ml. 02/gm./hr. at 30° C, when correctedto 20° C, on the assumption that the Qio is ca. 2.3, gives a value identical withthat which is found for winter R. temporia. April frogs, to be sure, would havea higher metabolic rate due to their feeding, increased physical activity and in-cipient spawning. Indeed, Krogh (1904) found the highest oxygen uptake occursduring the spawning season. Krogh was interested in another phase of frogmetabolism and, owing to experimental necessity, forcibly confined his animals.Even so, he obtained values of ca. 0.108 and 0.59-0.113 ml. O2/gm./hr. at 20° Cfor winter R. fusca and R. esculenta, respectively.It is interesting to note that Calhoon and Angerer (1955) using the sameapparatus, frog stock and experimental technique, except for the omission ofsnaking of the volumeters during the course of the experiment, (cf. their results,1953, p. 242) obtained a mean of 0.156 ml. O2/gm./hr. at 30° C. This may representa truer "basal" value for R. pipiens, though the value is approximately at thelimit of experimental error and its statistical significance is questioned.Renal damaged frogs. The daily mean values for the oxygen consumption of12 renal damaged frogs is presented in curve R, figure 1. After an initial decreaseof 6.8 percent at the end of the first postoperative day, the data show little vari-ation thereafter with time when compared with normal frogs (curve N). Thereis no statistical significance between these values (curve R) and those for normalfrogs (curve N). All subsequent comparisons, unless otherwise stated, are madewith unoperated frogs. These controls are used in place of renal damaged frogsin order that data from "paired" and "unpaired" adrenalectomized frogs maybe compared. There is little reason to believe that sham operations with theirconcomitant renal damage would alter the oxygen uptake of amphibious andaquatic animals as long as the regenerative area is relatively small compared tothe total respiring tissue mass. Furthermore, the amount of renal damage arising,of necessity, as a result of adrenal ablation, is apparently without injurious effectsto the frog (Angerer, 1950).Adrenalectomized frogs. Figure 1 gives data for the daily mean rates of oxygenconsumption for "paired" (curve A1) and "unpaired" (curve A) adrenalectomizedfrogs compared with their controls (curve N). While 71 percent of the "paired"frogs show a decrease in mean oxygen consumption at the end of the first post-operative day, a statistically significant decrease (ca. 13%, P<0.05) is observedby the end of the second postoperative day. Thereafter the values decreaseuntil the end of the fifth day when a mean of 0.095 ml./gm./hr. is obtained. Thisvalue is ca. 49 percent below that for the mean initial (zero) time or for the renaldamaged frogs at the end of a corresponding postoperative interval.A comparison of data for "paired" (curve A1) and "unpaired" (curve A)adrenalectomized frogs shows that both sets of data fall along essentially sigmoidcurves. Whether the curve form is significant, or fortuitous, is problematical.At best, it suggests many reactions are occurring simultaneously. To our know-ledge a comparable treatment of data for warm blooded animals does not exist.Secondly, there is a consistently greater decrease in mean oxygen uptake by theend of the fifth postoperative day for both winter (10%) and spring (12%) "paired"than for "unpaired" adrenalectomized frogs. Little variation is observed betweenNo. 5 OXYGEN CONSUMPTION OF FROGS 299winter (0.101 ml. 02/gm./hr. and spring 0.089 ml.) "unpaired" adrenalectomizedfrogs by the end of the fifth postoperative day. Thirdly, the death point for amajority {ca. 65%) of the adrenalectomized frogs under prevailing experimentalconditions occurs by the end of the fifth day, whereas normally it is between theseventh and twelfth postoperative day (Angerer, 1950). The last two items areprobably to be interpreted as due to the stress induced in handling and in sub-jecting the frogs to the relatively high environmental temperature. These twostressing agents would increase the demand for unavailable adrenocorticalhormone(s) (Hartman and Brownell, 1949). A decrease in metabolism followingadrenalectomy has been noted for several homoiotherms: cats (Aub, Formanand Bright, 1922), rabbits (Marine and Baumann, 1922), rats (Carr and Beck,1927), and so on. The chief difference is the extent of the metabolic depressionbetween homoiotherms {ca. 25%) and poikilotherms {ca. 50%).SUMMARYThe oxygen consumption of 20-30 gm. male frogs, R. pipiens, was studiedby direct-reading volumeters. All frogs studied may be placed in 3 major groups:A, unoperated (normal control, curve N, fig. 1); B, renal damaged (sham operatedcontrol curve R); C, "paired" (curve A1) and "unpaired" (curve A) adrenal-ectomized frogs (see '' method ").1. The mean oxygen consumption for winter and spring frogs under "basal"conditions is 0.175 ±0.003 ml./gm./hr. at 30° C; no statistical difference is ob-served between winter and spring stocks under the experimental conditionsemployed.2. There is no significant difference between the mean values for oxygenuptake for excessive renal damaged and normal frogs (cf. curves R and N).3. Adrenalectomized frogs show no significant change in oxygen consumptionat the end of the first postoperative day compared with unoperated or renaldamaged frogs. Thereafter, the means for both "paired" (curve A1) and "un-paired" (curve A) adrenalectomized frogs fall along a sigmoid curve (fig. 1).On the fifth postoperative day a mean of 0.095 ml. 02/gm./hr. is obtained. Thefigure represents a decrease of 46 percent below that of the controls.4. The stress of repeated handling and of subjecting adrenalectomized frogs torelatively high environmental temperatures depresses both their metabolism andviability (cf. curves A' and A).ACKNOWLEDGMENTSWe wish to thank Dr. Thomas B. Calhoon, formerly of this laboratory andnow of the Department of Physiology, Medical College of South Carolina,Charleston, for his help to M. C. M. in conjunction with the use of the respirometers.REFERENCESAngerer, C. A. 1940. The effect of thermal increments on and the subsequent adjustment ofthe protoplasmic viscosity of Amoeba proteus. Physiol. Zool. 13: 219-226.. 1942. Unpublished data.. 1950. Body weight, survival time, coloration, and water content of skeletal musclesof adrenalectomized frogs. Ohio Jour. Sci. 50: 103-106., and H. H. Angerer. 1949. The rate and total loss of body water on the survival timeof adrenalectomized frogs. Proc. Soc. Exper. Biol. and Med. 71: 661-665., and . 1950. Weight variations of adrenal ectomized frog muscles in differentconcentrations of Ringer's solutions. Proc. Soc. Exper. Biol. and Med. 73: 265-268.Aub., J. C, J. Forman, and E. M. Bright. 1922. The effect of adrenalectomy upon the totalmetabolism of the cat. Amer. Jour. Physiol. 61: 326-348.Calhoon, T. B., and C. A. Angerer. 1950. The effect of relatively large doses of sesame oilon oxygen consumption and the antagonistic action of adrenocortical extract. Endocrinol.46: 327-333.300 CLIFFORD A. ANGERER AND MARY C. MURRAY Vol. LV. 1951. Use of a new respirometer in studying the effects of adrenocortical extract ontotal metabolism. Fed. Proc. 10: 23.. 1953. A volumeter for respiration of aquatic animals. Ohio Jour. Sci. 53: 240-243.1955. Adrenal cortical extract on the oxygen consumption of normal frogs. Physiol.Zool. In press.Carr, C. J., and F. F. Beck. 1937. The metabolism of adrenalectomized rats. Amer. Jour.Physiol. 119: 589-592.Krogh, A. 1904. On the cutaneous and pulmonary respiration of the frog. (A contribution tothe theory of the gas exchange between the blood and the atmosphere.) Skand. Arch.Physiol. 15:328-419.Hartman, F. A., and K. A. Brownell. 1949. The adrenal gland. Lea and Febiger. Phil-adelphia.Loewy, 1926. Gas-und energie-wechsel. Tabulae Biologicae Periodicae 3: 504.Marine, D., and E. J. Baumann. 1922. VI. Further data on the effect of suprarenal in-sufficiency (by removal) in rabbits. Jour. Metabol. Research 2: 1-18.Williams, M. W., and C. A. Angerer. 1950. Adrenalectomy on frog skin potentials. Fed.Proc. 9:136.. 1952. Unpublished data.

Oxygen Consumption of Adrenalectomized and Renal Damaged Frogs

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Oxygen Consumption of Adrenalectomized and Renal Damaged Frogs

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