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Variations in Ascorbic Acid Concentrations in the Liver, Gonads, Kidney, and Blood Serum of Feral Blue Rock Pigeon (Columba livia gmelin) During the Breeding and Nonbreeding Seasons
PHYSIOLOGY AND REPRODUCTION Variations in Ascorbic Acid Concentrations in the Liver, Gonads, Kidney, and Blood Serum of Feral Blue Rock Pigeon (Columba livia gmelin) During the Breeding and Nonbreeding Seasons V. C. KOTAK' and P. M. AMBADKAR Division of Reproductive Physiology, Department of Zoology, The M. S. University of Baroda, Baroda 390 002, India (Received for publication November 15, 1983)
1985 Poultry Science 64:542-544 INTRODUCTION
Most invertebrates and fish lack the ability to synthesize ascorbic acid (AA) (Chatterjee, 1973). Phylogenetically, the capacity to synthesize AA starts with the amphibian kidney, resides in the reptilian kidney, and is transferred to the liver of mammals (Chatterjee, 1973). Because the enzymes for synthesizing AA are absent in the guinea pig, bat, monkey, and man, these animals depend on a dietary source for this vitamin. Biosynthesis of AA in nonpasserine birds occurs in the kidney, whereas in higher passerines, the kidney as well as the liver possess this capacity (Ray Chaudhri and Chatterjee, 1969; Chatterjee, 1973). The red-vented bulbul (Picnonotus cafer) does not possess the ability to synthesize AA (Chatterjee et al. (1968). Chinoy (1972a) has studied AA in different tissues of the pigeon (Columba livia) and is of the opinion that ascorbic acid participates in the general metabolic turnover by donating its free radicals. The influence of AA on biogenesis of hormones in the brain, adrenals, and testes has been investigated (Szent-Gyorgii, 1957; Biswas and Deb, 1970; Chinoy, 1970, 1972a,b). The interrelationship between AA and testosterone levels has been elucidated (Dieter, 1969;
1 Department of Biosciences, Sardar Patel University, Vallabh Vidyanagar 388120, India.
542
Majumdar and Chatterjee, 1974; Chinoy and Parmar, 1975; Chinoy et al, 1978). It is known that AA facilitates resistance to cold stress, accelerates haematopoiesis, and enhances the laying down of intercellular substances (Reid, 1954) — e.g., the synthesis and maturation of collagen, reticulin, dentin, and bone matrix — and to influence metabolism of glycosaminoglycans. The present investigation reports the concentrations of AA in the liver, gonads, kidney, and blood serum of feral pigeons during the breeding and nonbreeding phases of their annual reproductive cycle; the latter has been described previously (Ambadkar and Kotak, 1976a,b, 1977, 1978; Kotak, 1979, 1983). An attempt has been made to determine possible interrelationships between AA turnover and seasonal gonadal condition. MATERIALS AND METHODS
Both sexes of adult feral blue rock pigeons (Columba livia gmelin) were sacrificed with an air gun between 0900 and 1000 hr. Blood was collected from the jugular vein or heart. Birds were then immediately (within less than 5 min) brought to the laboratory. Part of the right liver lobe, first left kidney lobe, and a part of left gonad were used for biochemical measurements employing the method of Roe (1954). The AA was extracted with 6% TCA, which also reduces pH thus and prevents its catalytic oxidation.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
ABSTRACT To elucidate the possible involvement of ascorbic acid in gonadal steroidogenesis and role of gonadal steroids on the vitamin's turnover, ascorbic acid levels were quantified in the liver, gonads, kidney, and blood serum in botli sexes of the Indian feral blue rock pigeon (Columba livia gmelin) during the breeding (March to April) and nonbreeding (June to July) seasons. Low concentrations were observed during the breeding season and significantly higher cencentrations during the nonbreeding season. These changes are discussed in relation to the synthesis and distribution of ascorbic acid, gonadal steroidogenesis, and probable influence of gonadal function on ascorbic acid levels. (Key words: ascorbic acid, feral pigeon, seasonal, reproduction)
543
SEASONAL ASCORBIC ACID IN PIGEONS
The AA was oxidized to dehydroascorbic acid (DHA) by shaking the extract with activated charcoal. A 4-ml aliquot of the filterate was incubated with 2, 4-dinitrophenyl hydrazine for 3 hr at 37 C, thus yielding an osazone. This was allowed to react with 85% sulphuric acid, forming a reddish brown color, which was read colorimetrically at 540 m/i. Replicate assessments were carried out on all tissues. The breeding status of the birds was carefully ascertained and only adults were employed. RESULTS AND DISCUSSION
TABLE 1. Ascorbic acid levels (mean ± SD) in various organs and serum of the feral blue rock pigeon during the breeding (March to April) and nonbreeding (June to July) seasons1 Months
Liver
Kidney 2
Gonads (mg/100 g wet weig. i . . \ ,"t;
Serum 2 (mg/100 ml)
Males March April June
44.09 ± 4.51 3
{ 44.15 ± 3.40 55.29 ± 3.82
July
45.70 ± 4.12
{ 36.30 ± 5.08 98.35 + 10.31
19.07 + 4.39
5.23 ± .55 3.60 + .97
{ 34.43 ± 6.05
18.93 + 1.62
3.09 ± 1.18 4.14+ .52
{ 18.18 ± 2.14
{ 58.51 ± 4.59
{ 65.02 + 6.86
35.83 ± 3.68 38.26 ± 3.97
37.67 + 4.68 57.66 ± 6.21
16.40 + 3.20 14.43 + 2.73
3.13 ± .84 3.45 + .51
75.70 + 4.31
19.30 + 2.69 30.44 + 4.82
3.93 + .53 4.00 + .65
Females March April June
July
{
63.08 ± 5.10
{ 57.23 ± 5.31
{
{ 60.05 ± 2.98
1
Each mean value represents a minimum of 6 birds.
2
Unmarked (serum and female kidney) are nonsignificant: t < t „ .
3 Bracketed difference between the combined values of breeding (March-April) and non-breeding (JuneJuly) are highly significant as indicated by the Student's t test: P ( l t l ) > t 2 = .01.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
Among the three organs, the kidney had the lowest concentration of AA, being about 50% of the concentration found in the liver and gonads. Serum levels varied between 3.00 and 5.23 mg/100 ml. All the observed values are shown in Table 1. Synthesis and distribution of AA in cockerels are under hormonal control and the distribution of AA in pigeons is similar to that of the fowl (Dieter, 1969). The adrenal cortex and interstitial Leydig cells of birds and mammals have high AA content. The AA, monodehydro ascorbic acid (MDHA), and DHA are all involved in steroidogenesis in these sites (SzentGyorgii, 1957; Bacq and Alexander, 1961; Biswas and Deb, 1970; Chinoy, 1972a,b; Chinoy et al. 1978). In normal toads, administration of AA in conjunction with LH in-
creases the testicular A 5 -30-hydroxysteroid dehydrogenase (^5 -30-HSDH) activity, whereas hypophysectomy reduces the same (Biswas, 1969), which suggests that AA acts in conjunction with luteinizing hormone (LH) to augment testicular &5 -3/3-HSDH and thereby enhance steroidogenesis. In the current work, significantly high concentrations of hepatic, renal, and gonadal AA levels (except for the female kidney where the difference was statistically nonsignificant, see Table 1) were observed for the months of June and July (nonbreeding), a time when &5 -30-HSDH activity in the gonads is minimal (Ambadkar and Kotak, 1976b, 1977). An ultrastructural profile of the interstitial Leydig cells also reveals a nonsecretory phase during this season (Kotak, 1983), and gonadal weights are low (Kotak, 1979). Perhaps due to less LH, the steroid synthesizing enzyme had low activity. Significantly higher AA contents of liver, kidney, and gonads during the nonbreeding months could lead to decreased utilization and accumulation of AA in bound form. Many birds and mammals have bound AA in the liver and kidney (Needham, 1963; Malakar, 1963). Higher and significantly low AA levels in immature and laying birds, respectively, have been found in White Leghorn hens (Prabhakar et al. (1975). In contrast, the adminstration of testosterone propionate to intact 10-week-old cockerels significantly decreases renal glu-
KOTAK AND AMBADKAR
544
In summary, low levels of AA in the breeding season were possibly due to increased utilization, probably for steroidogenesis. Significantly higher concentrations of AA during the nonbreeding season were perhaps suggestive of unaltered synthesis coupled with postnuptial nonutilization of AA, which may result in its accumulation in the tissues. ACKNOWLEDGMENTS We are thankful to R. V. Shah (then Head of the Zoology Department, Baroda) for the facilities. We also appreciate the help of Subhash Bhatt, Mathematics Department, Sardar Patel University, Vallabh Vidyanagar. REFERENCES Ambadkar, P. M., and V. C. Kotak, 1976a. Seasonal variations of lipids and cholesterol in wild pigeon Columba livia: A quantitative study. J. Anim. Morphol. Physiol. 2 3 : 1 0 1 - 114. Ambadkar, P. M., and V. C. Kotak, 1976b. Observations on cyclic histochemical variations of lipids and steroid dehydrogenase in testes of feral rock pigeon Columba livia. Pavo 14: 80-87. Ambadkar, P. M., and V. C. Kotak, 1977. Seasonal variations of lipids and steroid dehydrogenases in the ovaries of feral blue rock pigeon Columba livia: A histochemical study. J. Anim. Morphol. Physiol. 24:118-126. Ambadkar, P. M., and V. C. Kotak, 1978. Histochemical observations on 3 OC -and 17/3-hydroxysteroid dehydrogenase in extra gonadal tissues of feral blue rock pigeon Columba livia G. Ind. J. Exp. Biol. 16:298-301. Bacq, Z. M., and P. Alexander, 1961. Fundamentals of Radiobiology. Pergamon Press, New York, NY. Biswas, N. M., 1969. A 5 -3/3-Hydroxysteroid dehydrogenase in toad testis. Synergistic action of ascorbic acid and luteinizing hormone. Endocrinology 85:981-983. Biswas, N. W., and C. Deb, 1970. In vitro studies on the effects of ascorbic acid biosynthesis in
animals. Ann. New York Acad. Sci. 92: 36—56. Chatterjee, I. B., Kar, N. C , and G. C. Chatterjee, 1968. Ascorbic acid requirement of the bird Pycnonotus cafer. Ind. J. Exp. Biol. 6:103— 105. Chatterjee, I. B., 1973. Evolution and biosynthesis of ascorbic acid. Science 182:1271-1272. Chinoy, N. J., 1970. Histochemical localization in animal tissues of a special peroxidase which generates the free radical of ascorbic acid. Stain Technol. 4 5 : 9 9 - 1 0 3 . Chinoy, N. J., 1972a. Ascorbic acid levels in avian tissues and its metabolic significance. Acta Zool. 53:121-126. Chinoy, N. J., 1972b. Ascorbic acid levels in mammalian tissues and its metabolic significance. Comp. Biochem. Physiol. 42:945. Chinoy, N. J., and P. Y. Parmar, 1975. Ascorbic acid metabolism in post-hatched chicken tissues under normal and cold-stress conditions. I. In some ascorbic acid and hormone synthesizing tissues. Pavo 13:54—65. Chinoy, N. J., L. Seethalaxmi, and Y. D. Singh, 1978. Beneficial role of ascorbic acid in vasectomy. Ind. J. Exp. Biol. 16:162-165. Dieter, M. P., 1969. Hormonal control of the synthesis and distribution of ascorbic acid in cockerel (Callus domesticus). Proc. Soc. Exp. Biol. Med. 130:210-213. Grollman, A. P., and A. L. Lehninger, 1957. Enzymic synthesis of L-ascorbic acid in different animal species. Arch. Biochem. Biophys. 69:458-467. Kotak, V. C , 1979. Certain histoenzymological and endocrinological observations on the reproductive cycles of the Indian feral blue rock pigeon Columba livia Gmelin. Ph.D. Thesis, Univ. Baroda, India. Kotak, V. C, 1983. Ultrastructure of the Leydig cells of feral pigeon during the non-breeding phase. Nucleus 26:86-89. Majumdar, P. K., and G. C. Chatterjee, 1974. Effects of testosterone and human chorionic gonadotrophin on the enzymes involved in the metabolism of L-ascorbic acid. Ind. J. Exp. Biol. 12:387-388. Malakar, M. C , 1963. Bound ascorbic acid in liver. Nature 198:185. Needham, J., 1963. Vitamins in ontogenesis. Chemical Embryology, Vol III. Hafner Publ. Co., New York, NY. Prabhakar, M. R., A. K. Chatterjee, and I. C. Datta, 1975. Influence of puberty and egg formation upon the cholesterol and ascorbic acid content of ovary and fallopian tube in White Leghorn birds. Indian J. Anim. Sci. 45: 284-286. Ray Chaudhri, C , and I. B. Chatterjee, 1969. Lascorbic acid in birds. Phylogenetic trends. Science 164:435-436. Reid, M. E., 1954. Ascorbic acid. VIII: Effects of deficiency in animals. The Vitamins. Academic Press, New York, NY. Roe, J. H., 1954. Chemical determination of ascorbic acid, dehydroascorbic acid and di-keto gulonic acids. Pages 115— 139 in Methods of Biochemical Analysis. David Glick, ed. Interscience Publ. Inc., New York, NY. Szent Gyorgii, A., 1957. Bioenergetics, Academic. Press, New York, NY.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
conate NADPH-oxidoreductase activity, and subsequently, hepatic and renal vitamin levels were elevated (Dieter, 1969). According to Grollman and Lehninger (1957), only the kidney contains all three enzymes for the vitamin synthesis in pigeons and chicken. In the present work, significantly lower AA in the liver, gonads, and kidney (except for the female kidney) during the breeding season may indicate an increased utilization. Blood serum of male birds registered higher values in March at a time when gonadal A 5 -3/3-HSDH activity was high and total cholesterol revealed depleted levels (Ambadkar and Kotak, 1976a,b, 1977). Subcellular images of the Leydig cells during these months indicated active androgen synthesis (unpublished observations).
1985 Poultry Science 64:542-544 INTRODUCTION
Most invertebrates and fish lack the ability to synthesize ascorbic acid (AA) (Chatterjee, 1973). Phylogenetically, the capacity to synthesize AA starts with the amphibian kidney, resides in the reptilian kidney, and is transferred to the liver of mammals (Chatterjee, 1973). Because the enzymes for synthesizing AA are absent in the guinea pig, bat, monkey, and man, these animals depend on a dietary source for this vitamin. Biosynthesis of AA in nonpasserine birds occurs in the kidney, whereas in higher passerines, the kidney as well as the liver possess this capacity (Ray Chaudhri and Chatterjee, 1969; Chatterjee, 1973). The red-vented bulbul (Picnonotus cafer) does not possess the ability to synthesize AA (Chatterjee et al. (1968). Chinoy (1972a) has studied AA in different tissues of the pigeon (Columba livia) and is of the opinion that ascorbic acid participates in the general metabolic turnover by donating its free radicals. The influence of AA on biogenesis of hormones in the brain, adrenals, and testes has been investigated (Szent-Gyorgii, 1957; Biswas and Deb, 1970; Chinoy, 1970, 1972a,b). The interrelationship between AA and testosterone levels has been elucidated (Dieter, 1969;
1 Department of Biosciences, Sardar Patel University, Vallabh Vidyanagar 388120, India.
542
Majumdar and Chatterjee, 1974; Chinoy and Parmar, 1975; Chinoy et al, 1978). It is known that AA facilitates resistance to cold stress, accelerates haematopoiesis, and enhances the laying down of intercellular substances (Reid, 1954) — e.g., the synthesis and maturation of collagen, reticulin, dentin, and bone matrix — and to influence metabolism of glycosaminoglycans. The present investigation reports the concentrations of AA in the liver, gonads, kidney, and blood serum of feral pigeons during the breeding and nonbreeding phases of their annual reproductive cycle; the latter has been described previously (Ambadkar and Kotak, 1976a,b, 1977, 1978; Kotak, 1979, 1983). An attempt has been made to determine possible interrelationships between AA turnover and seasonal gonadal condition. MATERIALS AND METHODS
Both sexes of adult feral blue rock pigeons (Columba livia gmelin) were sacrificed with an air gun between 0900 and 1000 hr. Blood was collected from the jugular vein or heart. Birds were then immediately (within less than 5 min) brought to the laboratory. Part of the right liver lobe, first left kidney lobe, and a part of left gonad were used for biochemical measurements employing the method of Roe (1954). The AA was extracted with 6% TCA, which also reduces pH thus and prevents its catalytic oxidation.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
ABSTRACT To elucidate the possible involvement of ascorbic acid in gonadal steroidogenesis and role of gonadal steroids on the vitamin's turnover, ascorbic acid levels were quantified in the liver, gonads, kidney, and blood serum in botli sexes of the Indian feral blue rock pigeon (Columba livia gmelin) during the breeding (March to April) and nonbreeding (June to July) seasons. Low concentrations were observed during the breeding season and significantly higher cencentrations during the nonbreeding season. These changes are discussed in relation to the synthesis and distribution of ascorbic acid, gonadal steroidogenesis, and probable influence of gonadal function on ascorbic acid levels. (Key words: ascorbic acid, feral pigeon, seasonal, reproduction)
543
SEASONAL ASCORBIC ACID IN PIGEONS
The AA was oxidized to dehydroascorbic acid (DHA) by shaking the extract with activated charcoal. A 4-ml aliquot of the filterate was incubated with 2, 4-dinitrophenyl hydrazine for 3 hr at 37 C, thus yielding an osazone. This was allowed to react with 85% sulphuric acid, forming a reddish brown color, which was read colorimetrically at 540 m/i. Replicate assessments were carried out on all tissues. The breeding status of the birds was carefully ascertained and only adults were employed. RESULTS AND DISCUSSION
TABLE 1. Ascorbic acid levels (mean ± SD) in various organs and serum of the feral blue rock pigeon during the breeding (March to April) and nonbreeding (June to July) seasons1 Months
Liver
Kidney 2
Gonads (mg/100 g wet weig. i . . \ ,"t;
Serum 2 (mg/100 ml)
Males March April June
44.09 ± 4.51 3
{ 44.15 ± 3.40 55.29 ± 3.82
July
45.70 ± 4.12
{ 36.30 ± 5.08 98.35 + 10.31
19.07 + 4.39
5.23 ± .55 3.60 + .97
{ 34.43 ± 6.05
18.93 + 1.62
3.09 ± 1.18 4.14+ .52
{ 18.18 ± 2.14
{ 58.51 ± 4.59
{ 65.02 + 6.86
35.83 ± 3.68 38.26 ± 3.97
37.67 + 4.68 57.66 ± 6.21
16.40 + 3.20 14.43 + 2.73
3.13 ± .84 3.45 + .51
75.70 + 4.31
19.30 + 2.69 30.44 + 4.82
3.93 + .53 4.00 + .65
Females March April June
July
{
63.08 ± 5.10
{ 57.23 ± 5.31
{
{ 60.05 ± 2.98
1
Each mean value represents a minimum of 6 birds.
2
Unmarked (serum and female kidney) are nonsignificant: t < t „ .
3 Bracketed difference between the combined values of breeding (March-April) and non-breeding (JuneJuly) are highly significant as indicated by the Student's t test: P ( l t l ) > t 2 = .01.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
Among the three organs, the kidney had the lowest concentration of AA, being about 50% of the concentration found in the liver and gonads. Serum levels varied between 3.00 and 5.23 mg/100 ml. All the observed values are shown in Table 1. Synthesis and distribution of AA in cockerels are under hormonal control and the distribution of AA in pigeons is similar to that of the fowl (Dieter, 1969). The adrenal cortex and interstitial Leydig cells of birds and mammals have high AA content. The AA, monodehydro ascorbic acid (MDHA), and DHA are all involved in steroidogenesis in these sites (SzentGyorgii, 1957; Bacq and Alexander, 1961; Biswas and Deb, 1970; Chinoy, 1972a,b; Chinoy et al. 1978). In normal toads, administration of AA in conjunction with LH in-
creases the testicular A 5 -30-hydroxysteroid dehydrogenase (^5 -30-HSDH) activity, whereas hypophysectomy reduces the same (Biswas, 1969), which suggests that AA acts in conjunction with luteinizing hormone (LH) to augment testicular &5 -3/3-HSDH and thereby enhance steroidogenesis. In the current work, significantly high concentrations of hepatic, renal, and gonadal AA levels (except for the female kidney where the difference was statistically nonsignificant, see Table 1) were observed for the months of June and July (nonbreeding), a time when &5 -30-HSDH activity in the gonads is minimal (Ambadkar and Kotak, 1976b, 1977). An ultrastructural profile of the interstitial Leydig cells also reveals a nonsecretory phase during this season (Kotak, 1983), and gonadal weights are low (Kotak, 1979). Perhaps due to less LH, the steroid synthesizing enzyme had low activity. Significantly higher AA contents of liver, kidney, and gonads during the nonbreeding months could lead to decreased utilization and accumulation of AA in bound form. Many birds and mammals have bound AA in the liver and kidney (Needham, 1963; Malakar, 1963). Higher and significantly low AA levels in immature and laying birds, respectively, have been found in White Leghorn hens (Prabhakar et al. (1975). In contrast, the adminstration of testosterone propionate to intact 10-week-old cockerels significantly decreases renal glu-
KOTAK AND AMBADKAR
544
In summary, low levels of AA in the breeding season were possibly due to increased utilization, probably for steroidogenesis. Significantly higher concentrations of AA during the nonbreeding season were perhaps suggestive of unaltered synthesis coupled with postnuptial nonutilization of AA, which may result in its accumulation in the tissues. ACKNOWLEDGMENTS We are thankful to R. V. Shah (then Head of the Zoology Department, Baroda) for the facilities. We also appreciate the help of Subhash Bhatt, Mathematics Department, Sardar Patel University, Vallabh Vidyanagar. REFERENCES Ambadkar, P. M., and V. C. Kotak, 1976a. Seasonal variations of lipids and cholesterol in wild pigeon Columba livia: A quantitative study. J. Anim. Morphol. Physiol. 2 3 : 1 0 1 - 114. Ambadkar, P. M., and V. C. Kotak, 1976b. Observations on cyclic histochemical variations of lipids and steroid dehydrogenase in testes of feral rock pigeon Columba livia. Pavo 14: 80-87. Ambadkar, P. M., and V. C. Kotak, 1977. Seasonal variations of lipids and steroid dehydrogenases in the ovaries of feral blue rock pigeon Columba livia: A histochemical study. J. Anim. Morphol. Physiol. 24:118-126. Ambadkar, P. M., and V. C. Kotak, 1978. Histochemical observations on 3 OC -and 17/3-hydroxysteroid dehydrogenase in extra gonadal tissues of feral blue rock pigeon Columba livia G. Ind. J. Exp. Biol. 16:298-301. Bacq, Z. M., and P. Alexander, 1961. Fundamentals of Radiobiology. Pergamon Press, New York, NY. Biswas, N. M., 1969. A 5 -3/3-Hydroxysteroid dehydrogenase in toad testis. Synergistic action of ascorbic acid and luteinizing hormone. Endocrinology 85:981-983. Biswas, N. W., and C. Deb, 1970. In vitro studies on the effects of ascorbic acid biosynthesis in
animals. Ann. New York Acad. Sci. 92: 36—56. Chatterjee, I. B., Kar, N. C , and G. C. Chatterjee, 1968. Ascorbic acid requirement of the bird Pycnonotus cafer. Ind. J. Exp. Biol. 6:103— 105. Chatterjee, I. B., 1973. Evolution and biosynthesis of ascorbic acid. Science 182:1271-1272. Chinoy, N. J., 1970. Histochemical localization in animal tissues of a special peroxidase which generates the free radical of ascorbic acid. Stain Technol. 4 5 : 9 9 - 1 0 3 . Chinoy, N. J., 1972a. Ascorbic acid levels in avian tissues and its metabolic significance. Acta Zool. 53:121-126. Chinoy, N. J., 1972b. Ascorbic acid levels in mammalian tissues and its metabolic significance. Comp. Biochem. Physiol. 42:945. Chinoy, N. J., and P. Y. Parmar, 1975. Ascorbic acid metabolism in post-hatched chicken tissues under normal and cold-stress conditions. I. In some ascorbic acid and hormone synthesizing tissues. Pavo 13:54—65. Chinoy, N. J., L. Seethalaxmi, and Y. D. Singh, 1978. Beneficial role of ascorbic acid in vasectomy. Ind. J. Exp. Biol. 16:162-165. Dieter, M. P., 1969. Hormonal control of the synthesis and distribution of ascorbic acid in cockerel (Callus domesticus). Proc. Soc. Exp. Biol. Med. 130:210-213. Grollman, A. P., and A. L. Lehninger, 1957. Enzymic synthesis of L-ascorbic acid in different animal species. Arch. Biochem. Biophys. 69:458-467. Kotak, V. C , 1979. Certain histoenzymological and endocrinological observations on the reproductive cycles of the Indian feral blue rock pigeon Columba livia Gmelin. Ph.D. Thesis, Univ. Baroda, India. Kotak, V. C, 1983. Ultrastructure of the Leydig cells of feral pigeon during the non-breeding phase. Nucleus 26:86-89. Majumdar, P. K., and G. C. Chatterjee, 1974. Effects of testosterone and human chorionic gonadotrophin on the enzymes involved in the metabolism of L-ascorbic acid. Ind. J. Exp. Biol. 12:387-388. Malakar, M. C , 1963. Bound ascorbic acid in liver. Nature 198:185. Needham, J., 1963. Vitamins in ontogenesis. Chemical Embryology, Vol III. Hafner Publ. Co., New York, NY. Prabhakar, M. R., A. K. Chatterjee, and I. C. Datta, 1975. Influence of puberty and egg formation upon the cholesterol and ascorbic acid content of ovary and fallopian tube in White Leghorn birds. Indian J. Anim. Sci. 45: 284-286. Ray Chaudhri, C , and I. B. Chatterjee, 1969. Lascorbic acid in birds. Phylogenetic trends. Science 164:435-436. Reid, M. E., 1954. Ascorbic acid. VIII: Effects of deficiency in animals. The Vitamins. Academic Press, New York, NY. Roe, J. H., 1954. Chemical determination of ascorbic acid, dehydroascorbic acid and di-keto gulonic acids. Pages 115— 139 in Methods of Biochemical Analysis. David Glick, ed. Interscience Publ. Inc., New York, NY. Szent Gyorgii, A., 1957. Bioenergetics, Academic. Press, New York, NY.
Downloaded from http://ps.oxfordjournals.org/ by guest on April 23, 2015
conate NADPH-oxidoreductase activity, and subsequently, hepatic and renal vitamin levels were elevated (Dieter, 1969). According to Grollman and Lehninger (1957), only the kidney contains all three enzymes for the vitamin synthesis in pigeons and chicken. In the present work, significantly lower AA in the liver, gonads, and kidney (except for the female kidney) during the breeding season may indicate an increased utilization. Blood serum of male birds registered higher values in March at a time when gonadal A 5 -3/3-HSDH activity was high and total cholesterol revealed depleted levels (Ambadkar and Kotak, 1976a,b, 1977). Subcellular images of the Leydig cells during these months indicated active androgen synthesis (unpublished observations).