Bulletin ILLINOIS [Ta.tii.]:*a.l Histo]:*3r Su-rvey SXJLLETIN ^(laid F. Labisky Vliam L. Anderson Nutritional Responses of Pheasants to Corn, with Special Reference to High-Lysine Corn / NATURAL HISTORY SURVEY NOV 14 1973 LIBRARY T«-: OF ILLINOIS )E|ARTMENT OF REGISTRATION AND EDUCATION fURAL HISTORY SURVEY DIVISION ^ANA, ILLINOIS ir THE LIBRARY QEWE Nn\J 7- 1Q70 VOLUME 31, ARTICLE ILLIN^OIS Satu]:*ail History Survey Nutritional Responses of Pheasants to Corn, with Special Reference to High-Lysine Corn onid F. Labisky lilam L. Anderson rMOF ILLINOIS EPRTMENT OF REGISTRATION AND EDUCATION AURAL HISTORY SURVEY DIVISION RANA, ILLINOIS VOLUME 31, ARTICLE JULY, 1973 STATE OF ILLINOIS DEPARTMENT OF REGISTRATION AND EDUCATION BOARD OF NATURAL RESOURCES AND CONSERVATION Dean Babrinoeb, Ph.D., Chairman: Thomas Park, Ph.D., Biology; L. L. Sloss. Ph.D., Geology; (Vacant), ChemiatTv: Robert H. Anderson, B.S.C.E., Engineering; Charles E. Olmsted, Ph.D., Forestry; W. L. EVERITT, E.E., Ph.D., Representing the President of the University of Illinois; Roger E. Bbyler, Ph.D., Representing the President of Southern Illinois University. NATURAL HISTORY SURVEY DIVISION. Urbana. Illinois j SCIENTIFIC AND TECHNICAL STAFF ' George Sprugel, Jr., Ph.D., Chief Alice K. Adams, Secretary to the Chief Section of Economic Entomology William H. Luckmann, Ph.D., Entomologist and Head Willis N. Bruce, Ph.D., Entomologist Wayne L. Howe, Ph.D., Entomologist Stevenson Moore, III, Ph.D., Entomologist, Exten- Howard B. Petty, Ph.D., Entomologist, Extension James E. Appleby, Ph.D., Associate Entomologist Edward J. Armbrust, Ph.D., Associate Entomologist Marcos Kocan, Ph.D., Associate Entomologist Joseph V. Maddox, Ph.D., Associate Entomologist Ronald H. Meyer, Ph.D., Associate Entomologist Robert D. Pauscii, Ph.D., Associate Entomologist Ralph E. Sechbiest, Ph.D., Associate Entomologist John K. Bouseman, M.S., Assistant Entomologist George L. Godfrey. 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Professor of Zoology and Director of Co- operative Wildlife Research. Southern Illinois University; Parasitology, Norman D. Levine, Ph.D., Profes- sor of Veterinary Parasitology, Veterinani Research, and Zoology and Director of the Center for Human Ecology, University of Illinois: Entomology, Robert L. Metcalf, Ph.D., Professor of Zoology and of En- tomology and Head of the Department of Zoology. University of Illinois; and Gilbert P, Waldbauer, Ph.D., Professor of Entomology, University of Illinois; Statistics. Horace W. Norton. Ph.D., Professor of Sta- (i«(ica( Design and Analysis. University of Illinois. Section of Administrative Services Robert O. Watson, B.S., Administrator and Head Supporting Services Vebnon F. Billman. Maintenance Supervisor WiLMA G. DiLLMAN, Property Control and Trust Accounts ROBEBT O. Ellis. Assistant for Operations Lloyd E. Huffman. Stockroom Manager J. William Lusk, Mailing and Distribution Services Melvin E. Schwartz, Financial Records James E. 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Assistant Technical CONTENTS Acknowledgments 87 Methods 88 Feeding Trial I : Juvenile Hens 88 Feeding Trial II : Adult Hens 88 Collection of Data 89 Analyses 90 Findings 90 Juvenile Hens 90 Body Weight Changes 90 Food Consumption 91 Digestibility Coefficients 91 Calorie Utilization 92 Protein Utilization 92 Lysine Utilization 93 Protein vs. Lysine 94 Glands and Organs 94 Fat and Fatty Acids 94 Adult Hens 97 Body Weight Changes 97 Digestibility Coefficients 98 Calorie Utilization 98 Protein Utilization 98 Lysine Utilization 98 Glands and Organs 99 Fat and Fatty Acids 100 Discussion 100 Summary 107 Literature Cited 109 Index Ill This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It is 7 contribution from the Section of Wildlife Research of the Illinois Natural History Survey. (60378—5M—7-73) Nutritional Responses of Pheasants to Corn, with Special Reference to High-Lysine Corn Ronald F. Labisky William L.Anderson IN LATE 1963, Purdue University sci- entists discovered, by amino acid analysis, that the endosperm of maize [Zea mays) kernels homozygous for the opaque-2 mu- tant contained about 70 percent more lysine than the endosperms of kernels of normal hybrids (Mertz et al. 1964: Mertz 1966:12). The endosperm of opaque-2 also contained greater amounts of tryp- tophan than that of normal corn ( Pickett 1966:19). Lysine and tntophan are among those amino acids that are dietary essentials for protein synthesis in many animals, including man. The proteins in endo- sperm of normal corn are of low biologi- cal quality. Thus the opaque-2 mutant, which alters the amino acid composition (particularly that of lysine, tryptophan, and leucine) of the maize endosperm, has offered the potential of a type of corn having exceptional nutritional values. The superior nutritional benefits of this modified-protein corn (hereinafter termed high-lysine corn) for growth have already been demonstrated in feeding experiments with rats (Mertz et al. 1965; Mcrtz 1966), swine (Pickett 1966: Jensen et al. 1967). chicks ( Rogler 1966 . and turkeys (.Adams & Rogler 1970). The nutritional potential of high-ly- sine corn has led to predictions that this corn may replace a substantial acreage of normal-corn hybrids produced in the Corn licit during the 1970"s. The esti- mated acreage of high-lysine corn plant- ed in the United States in 1972 was 80,000-100,000 acres (D. E. Alexander. Lfniversity of Illinois, personal communi- cation, January 12, 1973). Inasmuch as corn is important in the diet of many wild animals, the widespread use of high- lysine corn offers a potential nutritional benefit to wild birds and mammals. Corn figures more prominently in the diet of midwestern pheasants, particu- larly in fall and winter, than it does for most wildlife species (Korschgen 1964: 170, 173). To illustrate, during fall and winter, corn constitutes at least 80 percent (by weight) of the total food intake by pheasants in thriving populations in east- central Illinois (Anderson & Stewart 1969:261; R. F. Labisky, unpublished data). Yet despite the importance of corn to pheasants, little is known of its nutritional attributes for growth, main- tenance, or reproduction. Furthermore, juvenile hens, in contrast to adult hens, sufl'er a disproportionately high rate of nonhunting mortality between fall and winter in Illinois (R. F. Labisky, unpub- lished data) . That the onset of this mortality among juvenile hens coincides with that time of the year at \vhich waste corn from the harvest suddenly becomes abundantly available suggests a potential causal link between unbalanced nutrition and mortality. Hence the objectives of this study were to ascertain the physio- logical responses of juvenile hen pheas- ants in fall, and of adult hen pheasants in late %vinter and early spring, to ex- clusive diets of both normal corn and hiffh-lysine corn. ACKNOWLEDGMENTS .Acknowledgment is due the following members of the Department of Agrono- mv, L'nixersity of Illinois. Dr. D. E. .\lexander supplied the corns, provided their lysine and fatty acid profiles, and offered advice on various aspects of the 87 88 Illinois Natural History Survey Bulletin Vol. 31, Art. 3 study. Dr. T. R. Peck and G. G. Stone offered laboratory facilities for, and ma- terially aided in, the analyses of pheasant excreta for nitrogen. Dr. I. de la Roche analyzed the fat samples for determina- tion of fatty acids. Dr. C. M. Wilson analyzed the commercial ration for ami- no acids. Dr. B. G. Harmon, Department of Animal Science, University of Illinois, supervised the analyses of pheasant ex- creta for lysine. Dr. G. C. Sanderson, Illinois Natural History Survey, offered editorial sugges- tions during preparation of the paper, and O. F. Glissendorf edited the final manuscript. D. R. Vance and J. E. Mc- Clendon of the Survey assisted in various aspects of the experiment. Special thanks are due Drs. Alexander and Harmon, and Dr. J. E. Savage, De- partment of Poultry Science, University of Missouri, for critically reviewing the manuscript. METHODS FEEDING TRIAL I: JUVENILE HENS The 21 juvenile hens used in the ex- periment were obtained from the Illinois State Game Farm, Yorkville, in 1966. These hens, which had hatched on June 20, were transported to Urbana on Sep- tember 13. The hens were held in two wire-bottomed 3.0 x 3.9 x 1.8-meter pens and fed a commercial flight and maintenance chow (FMC) until October 3 when they were individually placed, by random assortment, in 70 x 60 x 34-cm cages. The cages had thin-walled fiber- glass sides, top, bottom, and rear, which prevented sight contact between birds. The birds were fed a diet of two-thirds FMC and one-third normal corn (whole kernels) for the period October 3-14 to acquaint them with corn, and then an exclusive diet of FMC for the period October 15-20. Inasmuch as 19 of the 21 hens post- ed gains in body weight between Octo- ber 10 and October 20, the feeding trial was begun on the latter date. Three groups of 7 hens each were randomly selected to be fed exclusive, unrestricted diets of FMC, normal hybrid corn (Pio- neer 3306), or high-lysine corn (Table 1 ) , and water ad libitum. The FMC was pressed into corn-sized pellets for the feeding trials (see Frontispiece). The experiment was terminated 8 weeks later, December 15. One hen from the group of hens fed normal corn died from an injury during the trial. FEEDING TRIAL II: ADULT HENS The 12 adult hens, 3 and 4 years old, used in the feeding trial were also of game-farm origin. These hens had been transported to Urbana as juveniles, and subsequently maintained in wire-bot- tomed outdoor pens, similar to those used to house the juveniles. On February 7, 1967, these hens were individually placed, by random assortment, in the same cages in which the juveniles of Trial I had been held. They had been fed an intro- ductory diet of one-half FMC, one- fourth normal corn, and one-fourth high- lysine corn for the period February 1-7. Because of their quick acceptance of com, they were returned to an exclusive FMC diet on February 8. All 12 hens posted gains in body weight during the interval of February 27-March 6; therefore, the feeding trial was begun on the latter date. Six hens were offered a diet of normal corn and six hens a diet of high-lysine corn (Table 1); both groups had unrestricted access to water. The food intake by adult hens was re- stricted to 200 g of corn per bird per week. The corn was provided in two 100-g lots, on the first and fourth days of each week. This limited offering of corn was judged to be about 60 percent of a normal weekly intake, and was in- tended to simulate the estimated poten- tial food intake of wild hens subjected to the rigors of late winter in Illinois. The experiment was terminated after 7 weeks, on April 24. July, 1973 Labisky & Anderson: Nutritional Responses of Pheasants Table 1.—Mean concentrations of calories, crude protein, lysine in protein, and selected min- erals in diets of a commercial flight and maintenance chow (FMC), of normal corn, and of high-lysine corn that were fed to hen pheasants in 1966 and 1967. 90 Illinois Natural History Survey Bulletin Vol. 31, Art. 3 recorded for kidneys and endocrine glands are for paired (right and left) measurements. ANALYSES Each sample of food and excreta was oven-dried at 60°C for 142 hours, finely ground, and then sealed in a sterile plas- tic bag for subsequent determination of nitrogen (crude protein), lysine, and caloric content. Nitrogen content of foods and excreta was determined by Kjeldahl procedures; the crude protein content of each item was calculated as nitrogen x 6.25. Crude protein determinations were made for five samples of each of the three foods, and for single samples of dried excreta from each juvenile hen for each of the last 5 weeks of the 8-week experiment and for each adult hen for Weeks 5 and 7 of the 7-week experiment. Lysine in the pheasant excreta was measured, following acid hydrolysis under vacuum for 16 hours, by chromatographic analysis (Beckman Amino Acid Analyzer, Model 120). Lysine determinations for excreta were made from composites of the weekly samples for Weeks 4-8 of the 8-week experiment for each juvenile hen, and for Weeks 5 and 7 of the 7-week experiment for each adult hen. The amount of lysine in the foods (defatted) was also measured by chromatographic analysis; approximately 80 percent of the nitrogen in the foods was recovered as amino acids. Caloric content of foods and excreta was measured by standard caloric-bomb techniques. Calories were measured from three samples of each of the three foods, and from a composite of the five and two weekly collections of excreta from each juvenile and each adult hen pheasant, respectively. The mineral content of the foods was derived by atomic absorption spectropho- tometry ( for Ca and Mg ) , flame spectro- photometry (for Na and K), and color- imetry (for P) . The null hypothesis, in all tests for de- termination of statistical differences, was accepted or rejected at the 0.05 level of probability. FINDINGS JUVENILE HENS Body Weight Changes The juvenile hens that were fed ex- clusive diets of FMC or high-lysine corn posted gains in body weight that aver- aged 98.4 and 23.4 g, respectively, during the 8-week feeding trial; those fed normal corn suffered losses that averaged 8.7 g ( Table 2 ) . Both groups of hens to which corn was fed exhibited marked declines in body weight during the first week of the feeding trial (Fig. 1). In the final analysis, all of the seven hens fed FMC, five of the seven hens fed high-lysine corn, and three of the six hens fed normal corn gained weight during the Table 2.—Body weight statistics for juvenile )ien pheasants fed exclusive diets of flight and maintenance chov/ (FMC), of normal corn, or of high-lysine corn for an 8-week period, October 20— December 15, 1966. Mean Body Weight (g) or Weight Change for Specified Diet FMC (n=7 Hens) Normal Corn (n=6 Hens) High-Lysine Corn (n=7 Hens) F Valuescjf) Initial weight (Oc- tober 20) Final weight (De- cember 15) Weight change 711.3 ± 29.2° 742.2 ± 36.9 809.7 ± 29.6 -1-98.4 ± 7.6 733.5 ± 35.8 -8.7 ± 14.7 776.1 -1-23.4 32.4 39.6 27.8 1.13:, 4.7O2. * Denotes statistical signtficancej P < 0.05. All combinations of paired means differed significantly. ° Standard errors. July, 1973 Labisky & Anderson: Nutritional Responses of Pheasants + 100- 91 •I (SlOg) (776fl) (OCTOBER 20) I DECEMBER 15)WEEKS OF STUDY Fig. 1.—Mean change in body weight (g) by weekly periods among juvenile hen pheasants fed exclusive diets of FMC, of normal corn, or of lysine corn; the vertical lines transecting the means ore standard errors. The mean initial and final body weights for each group are given at the left and right of the graph, respectively. Stotislicolly significant (P<0.05; 2 and 19 df) weight chonges occurred among diets for three of the eight successive weekly periods: initial to 1 (F=:3,73); 2 to 3 (F = 3.93); and 6 to 7 (F = 3.59). 8 weeks. The extremes in weight change among hens on each of the diets were: FMC, +128 and +59 g; high-lysine corn, +177 and -88 g; and normal corn, +44 and -71 g. Food Consumption The three diets, fed ad libitum, were consumed by the juvenile hens in signifi- cantly different amounts; the greatest in- take was of FMC and the lowest was of normal corn (Table 3). Inasmuch as the caloric, crude protein, and lysine con- tents of the high-lysine corn were either similar to or greater than those of normal corn (Table 1) , thereby discounting com- pensatory nutritional needs, the greater rate of consumption of high-lysine corn per hen suggested that it may have been more palatable^ to pheasants than nor- mal corn. Changes in body weight per 100 g of food consumed averaged +4.3, + 1.3, and -0.5 g on FMC, high-lysine corn, and normal corn, respectively. Diges+ibili+y Coefficients Significant differences in the digesti- bility coefficients- were exhibited by hens ^ Food intalte is oflcn depressed if the animal's diet is deficient in either protein or an indispensable amino add (see review by Harper, 1967). Therefore, the greater consumption of high-lysine corn over nor- mal corn by the hens may have reflected its higher lysine content rather than any superiority in palat- ability. - Digestibility coefficient = / Total dry weight of excreta VTotal dry weight of food con umed:t X 100 92 Illinois Natural History Survey Bulletin Vol. 31, Art. 3 on the different diets (Table 3). Hens fed FMC digested substantially less (59.1 percent) of tlieir ration than did those hens fed either normal corn (82.5 per- cent) or high-lysine corn (81.2 percent). In contrast to the corns, the compara- tively low rate of digestibility of FMC, in part, reflected its higher fiber content. Despite the similarity in the mean digesti- bility coefficients of the two corns, they were significantly different because of the extremely narrow range of variation in the digestibility of each of the corns by the individual hens. Calorie Ufilizafion Although the total intake of calories by the juvenile hens was significantly greater for those fed FMC than for those fed either normal corn or high-lysine corn, the utilization of the calories by the hens receiving the FMC was markedly less than for those receiving either of the corns; the metabolizabilitv coeffi- cients^ were 66.3, 84.8, and 83.3 for FMC, normal corn, and high-lysine corn, respectively (Table 4). Because of this different proportionate utilization of cal- ories, there \vas no significant difference in the total number of calories utilized per hen for birds on the three diets during the 8-week trial. Juvenile hens obtained 2,837, 3,945, and 3.808 kcal of metab- olizable energy per kg of FMC, normal corn, and high-lysine corn consumed, re- spectively. Protein Utilization The intake of crude protein by the juvenile hens was significantly different among the birds fed the three diets, being more than twice as great for FMC as for either normal or high-lysine corn (Ta- ble 3 ) . The high intake of crude protein by the hens fed FMC reflected not only i •'* Mctabolizability Total ca [- (- Table 3.—Comparative consumption and utilization of three foods—flight and maintenance chow (FMC), normal corn, and high-lysine corn—fed as exclusive diets to different groups of juvenile hen pheasants for an 8-week period, October 20-December 15, 1966. All values are expressed as dry weight. Mean Value per Hen per Week for Specified Diet FMC (n=7 Hens) Normal Corn High-Lysine Corn F Values«it) (n=6 Hens) (n=7 Hens) Food consumed (g) Crude protein consumed (g)' Lysine consumed (g)'' Excreta (g)'' Digestibility coefficient (percent)'' Crude protein in excreta (percent)'' Crude protein utilized (g)'' (percent)'' Change in body weight (g) 282.4 ± 12.9° 205.6 ± 10.8 234.6 64.4 3.2 115.5 1.0 0.06 2.1 24.7 0.8 36.0 1.3 0.01 0.6 27.5 1.3 44.2 17.0 0.7 0.03 1.0 65.11.,n«- 59.1 ± 0.3 39.3 ± 1.5 16.2 ± 1.8 26.0 ± 2.5 -1-12.3 82.5 ± 0.2 47.9 ± 0.9 6.5 ± 0.4 27.2 ± 1.6 — 1.1 81.2 ± 0.3 47.2 ± 0.7 5.7 0.4 21.0 -t-2.9 2,305 716. 2.266. 20. 24, 3. .5L.nc* .51=.,3C* .81.,S3* 67,.s, 92- M ' 24,.»3 lU.SS De al signific P < 0.05. yielded ni0.05; 1 and 10 df), Tissue July, 1973 Labisky & Anderson: Nutritional Responses of Pheasants corn and high-lysine corn, respectively; their final body weights averaged 200- 300 g below normal weights reported for adult hens at the initiation of egg-laying by Breitenbach (1963:26), Labisky & Jackson (1969:720), and Anderson (1972:459). Any meaningful discussion of the physiological responses of pheasants to diets of normal corn and high-lysine corn must be prefaced by some knowledge of the nutritional requirements of the spe- cies. Unfortunately, data regarding nu- tritional requirements of pheasants are scarce, particularly for subadult birds, and must be extrapolated from informa- tion available for poultry'. The National Research Council (1971:15-16) lists the minimum metabolizable energy (ME) requirements for domestic chickens (Cal- lus gallus) and turkeys (Mcleagris gallo- pavo) as 2,900-3,095 kcal per kg of food for 14- to 20-week-old birds, and as 2,850 kcal for mature breeders. Thus, the ME obtained by pheasants, both juveniles and adults, from the normal corn and high-l)'sine corn fed in this study well exceeded the energy levels required by poultry (Table 14). The FMC, however, only provided growing pheasants 2,837 kcal of ME per kg of ration, slightly less than the minimum ME required by grow- ing poultry. However, Barrett & Bailey (1972:14, 16-17) recently reported that breeding pheasants can perform satis- factorily on diets containing about 2,500 kcal of ME per kg of ration if the protein level is above 13 percent. The rates of metabolizability of both noiTnal corn and high-lysine corn by ju- venile and adult hen pheasants in this study paralleled closely the general 80 percent metabolizability rate of normal corn by chickens (Ewing 1963 : 83) . How- ever, among pheasants, the metaboliza- bility of normal corn was slightly greater than for the high-lysine corn regardless of whether the corns were fed ad libitum to juveniles or in restricted quantities to adults (Tables 3 and 10). Also, ju- veniles metabolized proportionately more of each of the two corns than did adults, a difTerence that may have reflected the relative demands of growth. Our study showed that although the juvenile hen pheasants utilized a similar Table 14.—Levels of protein, metabolizable energy, and selected omino ocids recommended for poultry feeds in relation to quantities supplied in diets fed to pheasonts during this study. All levels, except those for metabolizable energy, ore expressed as percentages of total diet. Dietary Requirements' 102 Illinois Natural History Survey Bulletin Vol. 31, Art. 3 number of calories on all three diets — FMC, normal corn, and high-lysine corn —the proportionate utilization of calor- ies was inversely associated with caloric consumption ( Tables 4 and 11). Barrett & Bailey (1972:20), however, reported that breeder pheasants compensated for foods with low ME levels by increasing food consumption, and consequently maintained reasonably similar levels of caloric intake on diets containing from 2,100 to 3,400 kcal of ME per kg. This type of compensatory action did not oc- cur among juvenile hen pheasants fed different diets in our study. The diets of these birds, however, were more variable in ME, protein content, and amino acid patterns than the rations fed by Barrett and Bailey, and hence are not totally comparable. In our study, the similarity in caloric utilization by juvenile pheasants among the diets characterized by different levels of ME was achieved not by compensatory caloric intake but by compensatory me- tabolizability. Furthermore, adult hens did not show a greater proportionate uti- lization of calories from the corn than did juveniles, even though they were fed a restricted ration and consumed fewer calories than they would have consumed on an ad libitum diet of corn. The National Research Council (1971 : 19) listed the dietary protein require- ments for starting and growing pheasants at 30 percent. Dale & DeWitt (1958: 292) reported that the growth rates of young pheasants, to 10 weeks of age, were less on diets containing 15, 18, and 22 percent protein than on diets contain- ing 28 percent protein. The Council (1971:15-16) also listed the protein re- quirements of the chicken as 20-23 per- cent for chicks, 12-16 percent for growing chickens, and 15 percent for laying (breeding) chickens; comparable levels for domestic turkeys were 28, 14-20, and 14 percent, respectively. The reported protein content in the diets of wild hen pheasants in the Midwest ranged season- ally from a minimum of about 12 percent to a maximum of about 19 percent (Korschgen 1964:169, 174) . Collectively, these findings suggest that the protein needs of pheasants are probably satisfac- torily met at levels of 16-20 percent for juveniles older than 14 weeks and 15 percent for adult breeders. Therefore, in this study, the dietary protein levels for pheasants were sufficient in the FMC (22.9 percent), but insufficient in both normal corn (12.0 percent) and high- lysine corn (11.7 percent). Whereas consumption of crude protein by the juvenile hen pheasants differed among birds fed FMC, normal corn, and high-lysine corn (Table 3), the propor- tionate utilization of the protein con- sumed, irrespective of amount, was simi- lar on all diets. Thus, among juvenile pheasants the total amount of protein utilized was related directly to the amount consumed—a situation opposite that for caloric utilization. Although adult hens consumed similar amounts of protein from the two corns, which were fed at a restricted rate, they utilized 41 percent less of the protein from high-lysine corn than from normal corn (Table 10). Thus, although both corns yielded dietary protein levels that were unsatisfactory to juveniles and adults, the pheasants still failed to utilize about three-fourths of all the protein they consumed in corn. Eleven of the 23 verified amino acids in proteins are essential to birds; that is, they cannot be sufficiently synthesized by the bird and must be taken in via the diet. Ewing (1963:201, 203) points out that arginine, lysine, methionine, cystine, and tryptophan are particularly important to birds because they are essential amino acids that are in critical demand during avian growth and development ; the other amino acids are either synthesized by the bird or are present in ample quantities in most foods. Important to the understanding of the amino acid complex is the fact that a deficiency of any essential amino acid will not only reduce the utilization of other July, 1973 Labisky & Anderson: Nutritional Responses of Pheasants 103 amino acids, but will also reduce the utili- zation of the entire diet. Thus, although this paper is concerned principally with the growth-associated amino acid, lysine, other amino acids that are potentially important to pheasants cannot be ignored. The FMC ration provided to pheasants in this study offered adequate quantities of protein and the essential amino acids for both subadult and adult birds (Table 14). The normal corn and high-lysine com diets, while providing minimal amounts of protein, did not supply ade- quate quantities of amino acids. The amino acids most lacking in the corns were lysine and methionine. High-lysine corn, however, offered an amino acid profile superior to that of normal corn, and the profiles of both corns were more aligned with the requirements of adult birds than of growing pheasants. The importance of lysine to growth was well illustrated in a study by Baldini et al. (1953:946-948). They demon- strated that young bobwhites (Colinus virginianus) , which reportedly required diets with 28 percent protein, could sur- vive and grow well on diets containing as little as 20 percent protein as long as the diets contained adequate amounts of lysine. In their experiments, the addi- tion of 0.3 percent lysine to a base diet of 20 percent protein and 1.0 percent lysine produced a ration with growth and sur\ival qualities for bobwhites that \vere equal to those provided by a diet contain- ing 28 percent protein and 1.0 percent lysine; thus, 0.3 percent lysine essentially replaced 8.0 percent crude protein. Such findings offer support for our conclusion that the amount of lysine utilized by ju- venile pheasants contributed more signif- icantly to their growth than did the amount of crude protein utilized (Table 6). As with crude protein, pheasants ex- hibited no compensatory utilization of lysine; the utilization of lysine \vas related i directly to its intake for both juvenile and adult birds (Tables 5 and 12). Perhaps the most interesting observation was the extremely high utilization (99.2 percent) of lysine from high-lysine corn by juvenile pheasants. This rate of utilization was not maintained by adult hens. The role of inadequate nutrition- quantitative or qualitative—as regards the physiology of stress in vertebrates is poor- ly understood. Among mammals, the term "stress" has become almost synony- mous with increased adrenocortical ac- tivity (see review by Christian 1963). Pre- sumably, some adverse stimulus triggers, via the hypothalamus, an increased re- lease of adrenocorticotropic hormone (ACTH), which in turn results in the increased production and secretion of corticosteroids from the adrenal corte.x that are necessary for maintaining physi- ological homeostasis under the given stress. (Prolonged exposure by the ani- mal to an adverse stimulus may result in e.xhaustion of the adrenal cortex, the subsequent failure of corticosteriod pro- duction, and finally death.) To produce the additional corticosteriods, the adrenal corte.x undergoes hyperplasia and hyper- trophy—hence, enlargement of the gland. Thus, enlarged adrenals have become generally recognized as clinical evidence of acute or chronic distress in mammals. Whether enlarged adrenals are a mea- sure of stress in birds is not clear. Like Christian & Davis (1966:11-13), who found a direct relationship between adrenal size (of mature females) and population density for vole ( Microtus pennsylvanicus) (Neave & Wright 1968: 634) reported a positive correlation be- tween adrenal-weight indices and popu- lation density for ruffed grouse (Bonasa umbellus) . Breitenbach et al. ( 1963 : 34) reported that the adrenals of adult hen pheasants that were restricted in their food intake (45 g per day) did not hy- pertrophy; however, the adrenals of in- dividual hens, in noticeably poor condi- tion, exhibited a marked increase in size. Also, Newlon et al. (1964:538-539) ob- served that the adrenal weights of bob- whites were greatest for birds fed those foods which yielded the poorest perform- Illinois Natural History Survey Bulletin Vol. 31, Art. 3 ance in maintaining body weight. These observations, coupled with our findings that the adrenal weights of hen pheasants were inversely related to changes in body weight (Table 7 and 13; Fig. 1-3), sug- gest to us that enlarged adrenals offer diagnostic symptoms of the stresses of inadequate nutrition in pheasants, and possibly other birds. The deposition and mobilization of depot fats are dynamic processes—even in an animal in reasonably stable energy balance (White et al. 1968:500). We found that depot fat, both strip and vis- ceral, was greatest for juvenile hen pheas- ants fed high-lysine corn, intermediate for those fed FMC, and least for those fed normal corn (Table 7). In contrast, adult hens, fed restricted but equal amounts of the two corns, accumulated greater fat deposits on a diet of normal corn than on a diet of high-lysine corn (Table 13). The fat deposits from these adult hens, irrespective of the type of corn diet, were many times smaller than those reported in spring for confined hens fed a high-protein ration ad libitum (Breitenbach 1963:32) or for wild hens (Anderson 1972:461). Breitenbach et al. (1963:34) presented evidence that the storage of fat may be stimulated by increased amounts of adrenocorticoster- oids. Hence, if the production of corti- costeroids paralleled increased adrenal size, as would be expected, fat deposits should have been related directly to adrenal size. We did not observe this relationship among the hens in this study. The birds' depot fats, which represent their largest reservoir of energy, were re- lated to body weight, and therefore in- versely reflected the hens' day-to-day energy demands. Depot fat consists chiefly of triglycer- ides; fatty acids, both saturated and un- saturated, are hydrolized from triglycer- ides via the action of the lipases. We found that the ratios of saturated to un- saturated fatty acids in the visceral fat from juvenile and adult hen pheasants fed normal corn and from juvenile hens fed high-lysine corn were about 27:73 (Table 8). Correspondingly, normal corn and high-lysine corn contained sat- urated to unsaturated fatty acid ratios of 14:86 and 19:81, respectively (Table 1). Hence, there was some disparity in the distribution of saturated and unsaturated fatty acids between the depot fat of pheasants and their corn diets. Although the distribution of fatty acids in the depot fat of herbivorous galliform birds gener- ally reflects the composition of the diet (Moss & Lough 1968:559; West & Meng 1968^:438), West & Meng (1968a: 539) have also provided evidence that, at least for the redpoll {Acanthis flammea) , en- vironmental conditions and the physio- logical state of the bird also influence the fatty acid composition of the visceral fat. Interestingly, the ratio of saturated to unsaturated fatty acids in the visceral fat of the adult hens fed the restricted intake of high-lysine corn was 23:77, which represented an increase in unsaturated fatty acids over the 27 : 73 ratio recorded for juvenile hens fed either high-lysine corn or normal corn ad libitum and for adult hens fed the restricted diet of nor- mal corn. The shift by adult hens fed high-lysine corn to a fatty acid composi- tion of depot fat that more closely re- flected that of their high-lysine corn diet is not surprising because White et al. (1968:499) reported that fatty acid pro- files of depot fat resemble the dietary profiles more closely when the depot fat is being depleted. The adult hens fed high-lysine corn at the restricted rate were decreasing in body weight, and therefore probably drawing on the stores of saturated fatty acids for reserve energy. Under these conditions the replacement fatty acids reflected the high proportion of unsaturates in the high-lysine corn diet. Plant seeds abound in unsaturated fat- ty acids. The principal unsaturated fat- ty acids in the corns were oleic and lino- leic. Correspondingly, the principal fatty acids in the visceral fat of pheasants were oleic and linoleic (Table 8) ; however, the samples of visceral fat contained pro- July, 1973 Labisky & Anderson: Nutritional Responses of Pheasants 105 portionally more oleic acid and consider- ably less linoleic acid than either of the corns. Linoleic was the principal fatty acid in the depot fats of the heather-eat- ing red grouse (Lagopus lagopus scoticus) (Moss & Lough 1968:560-561) and of the willow-eating willow ptarmigan {La- gopus lagopus alascensis) (West & Meng 1968^:438). However, as we found for the pheasant, Walker (1964:63-64) re- ported the predominant fatty acid in the depot fat of the seed-eating bobolink {Dolichonyx oryzivorus) to be oleic acid. Oleic acid and linoleic acid seem to be the principal unsaturated fatty acids that characterize the depot fats of granivorous and herbivorous birds, respectively. The parathyroid glands secrete a hor- mone that functions importantly as a regulator of calcium, and probably phos- phorus, metabolism; high and low levels of circulating calcium act on the glands to inhibit or stimulate, respectively, se- cretion of the parathyroid hormone (Geschwind 1961:434-436). Hypertro- phied parathyroid glands and reduced levels of blood calcium are characteristic responses of laying chickens to low-calci- um diets (Bloom et al. 1960:207). In our study, we fed juvenile hen pheasants diets that ranged from about 14,000 ppm calcium for FMC to about 40 ppm calci- um for the corns (Table 1); corns in general are notoriously low in calcium content. The parathyroids from these juvenile hens fed normal corn and high- lysine corn weighed at least twice as much as the glands from those fed FMC (Table 7), and nearly twice as much as the glands from their wild counterparts ( Anderson 1972: 485 ) . Furthermore, the adult hens fed exclusive diets of the corns during late winter and early spring (Ta- ble 13) had parathyroids substantially I larger than those reported by Anderson 1(1972:485) for wild adult hens at a 'comparable time of the year. The hyper- [trophied parathyroid glands from hen pheasants fed exclusive diets of calcium- 'deficient corns constituted strong clinical evidence that the birds were suffering from a negative calcium balance. The gonadal recrudescence among pheasants in spring is a response, medi- ated via the hypothalamo-hypophyseal axis, to increasing photoperiod (Bisson- nette & Csech 1936:106; Hiatt & Fisher 1947:538, 543; Greeley & Meyer 1953: 353-354) . In Illinois, complete gameto- genesis among hens, as evidenced by egg- laying, is attained between late March and mid-April (Labisky & Jackson 1966: 382; Labisky 1968:69; Labisky & Jack- son 1969:719). In our study, none of the adult hens fed corn diets restricted to an intake of 200 g per week had initi- ated egg-laying at the conclusion of the experiment on April 24. Furthermore, the reproductive tracts of these hens, when compared to hens on unrestricted diets or in the wild (Breitenbach et al. 1963:29; Anderson 1972:484) were se- verely underdeveloped physiologically for late April. The lag in ovarian and ovi- ducal development was more pronounced for hens fed high-lysine corn than for those fed normal corn (Table 13). The findings from this and previous investigations of confined pheasants (Ger- stell 1942:68; Kozlik 1949:62; Breiten- bach et al. 1963:27; Gates & Woehler 1968:240) have demonstrated that delays in egg-laying are related to poor physical condition in spring; Edwards et al. (1964:278) hypothesized a similar situa- tion for wild pheasants. Also, poor physi- cal condition, usually the result of mal- nutrition, signifies reduced reserves of energy. Fisher ( 1967 : 121) cited evidence to show that domestic hens would cease egg production as soon as their protein reserves were exhausted after being- placed on a protein- or amino acid-defi- cient diet. Corns do not abound in pro- tein, and are deficient in one or more of the essential amino acids. The inhibitory effects of inadequate nutrition on reproduction in galliform birds, however, seem to be mediated through the hypothalamo-hypophyseal axis and not directly by protein or amino acid imbalances. Morris & Nal- bandov (1961:687) demonstrated that undernourished or starved domestic pul- 106 Illinois Natural History Survey Bulletin Vol. 31, Art. 3 lets failed to produce sufficient gonado- tropins to promote functional ovarian and oviducal development. Gates & Woehler ( 1 968 : 243 ) sum it up aptly : "It would appear that restoration of body condition [in pheasants] ... is a requisite for re- crudescence, normal rates of egg laying being delayed until energy is available for reproduction and the secretory integ- rity of the pituitary is restored." The nutritional responses of pheasants to normal corn and high-lysine corn are not consistent for birds of different ages. The weight gains and physiological par- ameters for juvenile hen pheasants fed high-lysine corn in autumn were distinct- ly more favorable than for those corres- pondingly fed normal corn. Yet adult hens fared far better on normal corn as an emergency food in late winter than they did on high-lysine corn. Thus, al- though potentially of benefit to growing pheasants, high-lysine corn may well be a detriment to mature pheasants. These conflicting results are at least partially explainable. The greater nutritional val- ue to young pheasants of high-lysine corn, when contrasted with normal corn, un- doubtedly lies in its higher content of the growth-oriented amino acid, lysine. Our findings indicated strongly that the weight gains of juvenile pheasants were dependent directly on the amount of ly- sine utilized. Evidence in support of these findings is provided by the research of Cromwell et al. (1967:711), who studied the nutritional responses of do- mestic chicks to the two corns: ". . . the beneficial effects of opaque-2 corn over normal corn appeared to be medi- ated solely through its higher lysine con- tent." These workers, however, fed the corns as a part of nutritionally balanced basal diets, and not as exclusive food items. They also pointed out (p. 712) that the beneficial responses exhibited by young animals to opaque-2 corn would probably be much less for mature animals that have a lower protein requirement. Furthermore, the probability of selecting genetically for a pheasant that responds to higher than normal dietary levels of ly- sine by exhibiting an improved rate of growth seems remote, as Godfrey (1968: 1565) found that the heritability for ly- sine utilization among Japanese quail [Coturnix coturnix japonica) was very low. Thus opaque-2 corn apparently is not a panacea for assuring adequate nu- trition in pheasants. Another modified- protein corn called floury-2, which con- tains higher concentrations of both lysine and methionine than those in most other corns (Nelson et al. 1965:1470; Crom- well et al. 1968 : 846) , may offer nutrition- al benefits for birds that are potentially superior to those provided by either normal corn or opaque-2 corn. Corn, an important food source to many species of wildlife, is a nutritive sta- ple for pheasants. To illustrate, Newlon et al. (1964:536-537), in evaluating foods for sustaining bobwhites, reported that the mean survival duration for ju- venile and adult birds fed an exclusive, ad libitum diet of normal corn in No- vember was 22.6 days; no bobwhite sur- vived the 38-day feeding trial. As ob- served in this study, however, exclusive ad libitum diets of normal corn sustained juvenile pheasants, with only minor weight losses, for 8 weeks in autumn with- out any mortality. Also, adult hens fed a restricted intake of normal corn (200 g per week) in late winter and early spring maintained their body weights for a 7-week feeding trial. If, however, pheasants are to parallel the annual cycle of body weights that normally character- ize thriving populations in the wild they must supplement their corn-dominated diets with food items more nutritiously balanced than high-lysine or normal corn. A plausible hypothesis, therefore, is that the high rates of nonhunting mortality observed among wild juvenile pheasants in Illinois during autumn (R. F. Labisky, unpublished data) may be directly or in- directly related to nutritive imbalances resulting from the surging availability of waste corn in the birds' diet. July, 1973 Labisky & Anderson : Nutritional Responses of Pheasants 107 SUMMARY Corn is an important food for many \vild animals and is especially prominent in the diet of wild pheasants in the United States. Yet despite its importance as a staple food of pheasants, knowledge of its nutritional value to the species is still quite limited. In 1963, scientists added another dimension to corn nutrition by discovering a modified-protein corn, opaque-2 corn (herein called high-lysine corn), which has substantially greater amounts of lysine in its endosperm than does normal corn. Lysine is one of the essential growth-promoting amino acids. The objectives of this investigation were to ascertain the physiological re- sponses of juvenile hen pheasants in au- tumn, and of adult hens in late winter and early spring to exclusive diets of normal corn and high-lysine corn. In the 8-week feeding trial for juveniles, October 20-December 15, 1966, 21 hens in three groups of 7 each were fed ex- clusive, ad libitum, diets of a balanced ration (FMG), normal corn, or high- lysine corn. Analyses of the three foods yielded the following: FMC-4.28 kcal per g, 22.8 percent protein, and 4.9 per- cent lysine in protein; normal corn—4.65 kcal per g, 12.0 percent protein, and 3.2 percent lysine in protein; and high-lysine corn—4.54 kcal per g, 11.7 percent pro- tein, and 4.7 percent lysine in protein. The feeding trial for juveniles coincided with the season in which juvenile hens, in contrast to adult hens, suffer dispro- portionately high rates of nonhunting mortality in Illinois, and also simultane- ously with the time that waste corn from the harvest suddenly becomes an abun- dant food source. Hence this phase of the study was designed partially to de- termine if juvenile mortality among wild pheasants was related to unbalanced nu- trition. In the 7-week feeding trial for adult hens, March 6-April 24,' 1967, 12 hens in two groups of 6 each were fed e.xclusive diets of normal corn or high- lysine corn at the restricted rate of 200 g per bird per week. This restricted in- take of food was intended to simulate the conditions that wild hens in the Midwest often confront in late winter and early spring. The juvenile hen pheasants fed ex- clusive diets of FMC and high-lysine corn for the 8-week period in autumn posted gains in body weight that aver- aged 98.4 g (13.8 percent) and 23.4 g (3.1 percent), respectively; correspond- ingly those juvenile hens fed normal corn suffered average losses of 8.7 g (1.2 per- cent). Wild juvenile hens averaged gains of 110 g (13.2 percent) for the comparable autumn period. Adult hens, each fed a restricted in- take of 200 g of corn per week for the 7-week period in late winter and early spring, averaged losses in body weight of 5.8 g (0.6 percent) for normal corn and 65.5 g (7.3 percent) for high-lysine corn. Whereas hen pheasants usually exhibit gains in body weight in late winter, reaching their maximum weight just prior to the onset of egg-laying (usually in April), the adult hens on the restrict- ed intake of corn averaged 200-300 g below the normal body weights of wild hens in April. The kcal of energy metabolized per kg of food consumed by juvenile hens was 2,837 (66.3 percent efficiency) for FMC, 3,945 (84.8 percent) for normal corn, and 3,808 (83.3 percent) for high-lysine corn; adult hens metabolized 3,921 kcal per kg (84.3 percent) of normal corn and 3,617 kcal per kg (79.6 percent) of high-l)sine corn. Juvenile hens, despite the differences in the yield of metaboli- zable energy among the foods, utilized the same number of calories on all three diets; the similarity in caloric utilization was achieved by compensatory metaboliz- ability and not by compensatory calor- ic intake. Unlike the situation for calories, hen pheasants exhibited no compensatory utilization of either crude protein or ly- sine. The total amounts of both protein and lysine utilized by hen pheasants Illinois Natural History Survey Bulletin Vol. 31, Art. 3 were related directly to dietary intake. Even though the dietary protein levels of both corns were unsatisfactory, the birds utilized only about one-fourth of the protein they consumed. The propor- tionate utilization of lysine by either adult or juvenile hens exceeded 85 per- cent for all diets. Interestingly, juvenile hens utilized 99 percent of the lysine consumed from high-lysine corn—a rate not achieved by adult hens. Most im- portant was the finding that the body weights of juvenile pheasants in autumn were more dependent on the amount of lysine utilized than on the amount of crude protein utilized. The adrenal weights of hen pheasants were inversely related to changes in body weight, which in turn was a reflection of the consumption of diets of different nutritional offerings. These findings in- dicated that enlarged adrenal glands in pheasants, and possibly other avian species, may offer diagnostic evidence for detecting the physiological stresses of un- balanced or inadequate nutrition. Depot fats of hen pheasants were di- rectly related to changing body weight, a relationship that reflected changing demands for energy for growth or main- tenance. The ratio of saturated : unsatur- ated fatty acids in the visceral fat of hens fed corn was about 1 : 3, which ^vas greater than that found in the corns. Oleic and linoleic were the principal fatty acids in corn, and, correspondingly, the most prominent in the depot fats of pheasants. Hypertrophied parathyroid glands from hen pheasants fed exclusive diets of calcium-deficient corns offered strong clinical evidence that wild hen pheasants on corn-dominated diets would suffer from a negative calcium balance. The reproductive tracts of adult hen pheasants fed restricted diets of normal corn or high-lysine corn, unlike those of wild hens or hens fed unrestricted diets, were severely underdeveloped in late April. The lag in ovarian and oviducal development was substantially greater for hens fed high-lysine corn than for those hens fed normal corn. The effects of inadequate nutrition are apparently mediated through the hypothalamo-hy- pophyseal-gonadal axis. The nutritional responses of young and adult hen pheasants to normal corn and high-lysine corn were not similar. The physiological profiles of juvenile hens fed high-lysine corn in autumn were distinctly more favorable than of those fed normal corn. The greater nutritional value to young pheasants of high-lysine corn, in contrast to normal corn, very likely is associated with its higher content of lysine, an essential growth-promoting amino acid. However, as an emergency food for adult hens in late winter or early spring, normal corn proved superior to high-lysine corn. 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Corn Industries Research Foundation, Com Refiners Association, Inc., Washington, D.C. 186 p. Rogler, John C. 1966. A comparison of I opaque-2 and normal corn for the chick, p. 23-25. In Edwin T. Mertz and Oliver E. Nelson (Editors), Proceedings of the high lysine corn conference. Corn Indus- tries Research Foundation, Corn Refiners ! Association, Inc., Washington, D.C. 186 p. Walker, Alma Toers. 1964. Major fatty i acids in migratory bird fat. Physiological Zoology 37(l):57-64. West, George C, and Martha S. Meng. 1968a. The effect of diet and captivity on the fatty acid composition of redpoll {Acan- this jlammea) depot fats. Comparative i Biochemistry and Physiology 25(2): 535- 540. , and . 1968fc. Seasonal changes i in body weight and fat and the relation of I fatty acid composition to diet in the willow i ptarmigan. Wilson Bulletin 80(4) : 426- 441. White, Abraham, Philip HandleRj and : Emil L. Smith. 1968. Principles of bio- - chemistry. Fourth edition. The Blakiston ' Division, McGraw-Hill Book Company, Inc., New York. 1187 p. INDEX Acanthis fiammea (i^e redpoll) Adrenal glands as indicator of stress, 103, 108 mean weights of, 95, 100 weight correlated with body weight, 94, 96 Adrenocortical activity, 103 Adrenocorticosteroids, 104 Adrenocorticotropic hormone (ACTH), 103 Amino acids analyzer, 90 requirements for, 101 essentia! to birds, 102 Arachidonic acid, 96 Argmine. 101, 102 Atomic absorption spectrophotometry, 90 B Bobolink, 105 Bobwhite. 103, 106 Body weight changes in, 90-91, 97, 107 correlated with adrenal weight, 94, 96 correlated with lysine utilization, 93—94 correlated with protein utilization, 93, 94 of wild pheasants, 100 procedures for recording, 89 Bonasa umbellus (see ruffed grouse) Calcium, 89, 90, 105. 108 Calories consumption of, 93, 98, 99 determination of, 90 in experimental diets, 89, 107 utilization of, 92. 93, 98. 99 Care and housing of pheasants, 88 Chicken, 87, 101, 102, 105, 106 Colinus iirginianus {see bobwhite) Colorimetr\-, 90 Coracobrachialis. 89 Corn as food for wild animals, 87, 107 chemical composition of, 89, 101 metabolizable energy (ME) of, 101 Pioneer 3306. 88, 89 Corticosteroids, 103, 104 Coturnix coturnix japonica {see Japanese quail) Cystine. 101, 102 Egg-laying timing of, 105 Excreta mean weights of, 92, 98 procedures for collecting, 89 Fat deposits deposition and mobilization, 104 mean weights of, 95, 100 Fat strip {see fat deposits) Fatty acids in experimental diets, 89 in pheasants, 94, 96, 100 saturated: unsaturated ratios, 89, 96, 100, 104, 108 Feeding trials procedures for, 88 Flame spectrophotometry, 90 Flight and maintenance chow (FMC) chemical composition of. 89, 101 Floury-2, 106 Food consumption by experimental hens, 91, 92, 98 procedures for measuring, 89 Callus gallus {see chicken) Gizzard, 95 Glands, 89-90, 94, 95, 96, 99, 100, 103-104, 105. 108 Gonadal recrudescence, 105, 106 H High-lysine corn defined, 87 Hypothalamo-hypophyseal axis, 105, 108 Hypothalamus, 103 Japanese quail, 106 Kidneys, 95 Kjeldahl procedures, 90 Diets chemical composition of, 89, 101 Digestibility coefficients defined. 91 for experimental diets, 91-92. 98 Dissecting procedures, 89 Dolichonyx oryzivorus {see bobolink) Lagopus lagopus alascensis (see willow- ptarmigan) Lagopus lagopus scoticus (see red grouse) Laurie acid, 96 Leucine. 87 Linoleic acid. 96. 104-105, 108 111 112 Illinois Natural History Survey Bulletin Vol. 31, Art. .3 L (cont.) Linolenic acid, 96 Liver, 95, 100 Lysine consumption of, 92, 93, 99 correlated with body weight, 93-94 determination of, 90 importance of in nutrition, 87, 103, 108 in experimental diets, 89, 101, 107 requirements for, 101, 102 utilization of, 93, 99 M Magnesium, 89, 90 Meleagris gallopavo {see turkey) Metabolizability coefficient defined, 92 for experimental diets, 92, 93, 98, 99, 107 Metabolizable energy (ME) requirements for, 101 supplied by experimental diets, 101 Methionine, 101, 102, 103 Microtus pennsylvanicus [see vole) Minerals, 89, 90, 105, 108 MortaUty related to diet, 106, 108 Muscles (see sternal muscles) Myristic acid, 96 N Nitrogen determination of, 90 Null hypothesis level accepted or rejected, 90 Parathyroid glands as indicator of negative calcium balance, 105, 108 functions of, 105 mean weights of, 95, 100 Parathyroid hormone, 105 Pectoralis thoracia, 89 Phosphorus, 89, 90 Potassium, 89, 90 Protein consumption of, 92, 98 correlated with body weight, 93, 94 determination of, 90 in diets of wild pheasants, 102 in excreta, 92, 98 in experimental diets, 89, 107 requirements for, 101, 102 utilization, 92-93, 98, 102 Rats, 87 Red grouse, 105 Redpoll, 104 Ruffed grouse, 103 Sodium, 89, 90 Spleen, 95 Stearic acid, 96 Sternal muscles, 95, 100 Stress, 103 Supracoracoideus ventral head of, 89 Swine, 87 Oleic acid, 96, 104-105, 108 Opaque-2, 87, 106, 107 Organs, 89-90, 94, 95, 99, 100 Ovary, 100 Oviduct, 100 Ovum, 100 Thymus glands, 85 Thyroid glands, 95, 100 Triglycerides, 104 Tryptophan, 87, 101, 102 Turkey, 87, 101, 102 Visceral fat {see fat deposits) Vole, 103 Palmitic acid, 96 Palmitoleic acid, 96 Pancreas, 95 w Willow ptarmigan, 105 Some Publications of the ILLINOIS NATURAL HISTORY SURVEY BULLETIN Volume 30, Article 3.-MigrationaI Behavior of Mallards and Black Ducks as Determined from Banding. By Frank C. Bellrose and Robert D. Crompton. September, 1970. 68 p., frontis., 25 fig., bibliogr., index. Volume 30, Article 4.-Fertilization of Estab- lished Trees: A Report of Field Studies. By Dan Neely, E. B. Hiraelick, and Webster R. Crowley, Jr. September, 1970. 32 p., fron- tis., 8 fig., bibliogr., index. Volume 30, Article 5.-A Survey of the Mussels (Unionacea) of the IlHnois River: A Pollut- ed Stream. By William C. Starrett. February, tis., 8 fig., bibliogr., index. Volume 30, Article 6.-Comparative Uptake and Biodegradability of DDT and Methoxy- chlor by Aquatic Organisms. By Keturah A. Reinbold, Inder P. Kapoor, William F. Childers, Willis N. Bruce, and Robert L. Metcalf. June, 1971. 12 p., frontis., 5 fig., bibliogr., index. Volume 30, Article 7.-A Comparative Study of Two Components of the Poinsettia Root Rot Complex. By Robert S. Perry. August, 1971. 35 p., frontis., 10 fig., bibliogr., index. Volume 30, Article 8.-Dynamics of Condition Parameters and Organ Measurements in Pheasants. By William L. Anderson. July, 1972. 44 p., frontis., 6 fig., bibliogr., index. Volume 31, Article l.-The Effects of Supple- mental Feeding and Fall Drawdowns on the Largemouth Bass and Bluegills at Ridge Lake, Illinois. By George W. Bennett, H. Wickliffe Adkins, and William F. Chil- ders. January, 1973. 28 p., frontis., 8 fig., bibilogr., index. BIOLOGICAL NOTES 71.-A Synopsis of Common and Economic Illinois Ants, with Keys to the Genera (Hymenoptera, Formicidae). By Herbert H. Ross, George L. Rotramel, and Wallace E. LaBerge. January, 1971. 22 p., 27 fig., bibliogr. 72.-The Use of Factor Analysis in Modeling Natural Communities of Plants and Ani- mals. By Robert W. Poole. February, 1971. 14 p., 14 fig., bibliogr. 73.-A Distributional Atlas of Upper Mississip- pi River Fishes. By Philip W. Smith, Alvin C. Lopinot, and William L. Pflieger. May, 1971. 20 p., 2 fig., 107 maps, bibliogr. 74.-The Life History of the Slenderhead Dart- er, Percina phoxocephala, in the Embarras River, Illinois. By Lawrence M. Page and Philip W. Smith. July, 1971. 14 p., 10 fig., bibliogr. 75.-Illinois Birds: Turdidae. By Richard R. Graber, Jean W. Graber, and Ethelyn L. Kirk. November, 1971. 44 p., 40 fig., bib- liogr. 76.-Illinois Streams: A Classification Based on Their Fishes and an Analysis of Factors Re- sponsible for Disappearance of Native Spe- cies. By Philip W. Smith. November, 1971. 14 p., 26 fig., bibliogr. 7 7.-The Literature of Arthropods Associated with Soybeans. I. A Bibliography of the Mexican Bean Beetle, Epilachna varivestis Mulsant (Coleoptera: Coccinellidae). By M. P. Nichols and M. Kogan. February, 1972. 20 p., 1 fig., bibliogr. 78.-The Literature of Arthropods Associated with Soybeans. II. A Bibliography of the Southern Green Stink Bug, Nezara viridula (Linneaus) (Hemiptera: Pentatomidae). By N. B. DeWitt and G. L. Godfrey. March, 1972. 23 p., 1 fig., bibliogr. 79.-Combined Culture of Channel Catfish and Golden Shiners in Wading Pools. By D. Homer Buck, Richard J. Baur, Charles F. Thoits III, and C. Russell Rose. April, 1972. 12 p., 3 fig., bibliogr. 80.-Illinois Birds: Hirundinidae. By Richard R. Graber, Jean W. Graber, and Ethelyn L. Kirk. August, 1972. 36 p., 30 fig., bibliogr. 81.-Annotated Checklist of the Butterflies of Illinois. By Roderick R. Irwin and John C. Downey. May, 1973. 60 p., 3 fig., 98 maps, bibliogr. 82.-Lactate Dehydrogenase Isozymes of Dar- ters and the Inclusiveness of the Genuj Percina. By Lawrence M. Page and Greg- ory S. Whitt. May, 1973. 7 p., 5 fig., bibliogr. CIRCULAR 46.-Illinois Trees: Their Diseases. By J. Ced- ric Carter. June, 1964. (Third printing, with alterations.) 96 p., frontis., 89 fig. 49.-The Dunesland Heritage of Illinois. By Herbert H. Ross (in cooperation with Illinois Department of Conservation). August, 1963. 28 p., frontis., 16 fig., bibliogr. 51.-Illinois Trees: Selection, Planting, and Care. By J. Cedric Carter. August, 1966. 123 p., frontis., 108 fig. 52.-Fertilizing and Watering Trees. By Dan Neely and E. B. Himelick. December, 1971. (Third printing.) 20 p., 9 fig., bibliogr. 53.-Dutch Elm Disease in Illinois. By J. Cedric Carter. October, 1967. 19 p., frontis., 17 fig. Li^t of available publications mailed on request No charge is made for publications of the Illinois Natural History Survey. A single copy of most publications will be sent free to anyone requesting it until the supply becomes low. Costly publications, more than one copy of a publication, and publications in short supply are subject* for special correspondence. Such correspondence should identify the writer and explain the use to be made of the publication or publications. Address orders and correspondence to the Chief,