The Influence of Photoperiod on Body Weight Gain, Body Composition, Nutrient Intake and Hormone Secretion H.

Allen Tucker, D. Petitclerc and S. A. Zinn J Anim Sci 1984. 59:1610-1620.

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THE INFLUENCE OF PHOTOPERIOD ON BODY WEIGHT GAIN, BODY COMPOSITION, NUTRIENT INTAKE AND HORMONE SECRETION 1'2
H. Allen Tucker, D. Petitclerc s and S. A. Zinn Michigan State University 4, East Lansing 4.8824-1225

Summary

Increasing daily light exposure from 8 to 16 h increases average daily body weight gains of sheep and Holstein cattle but reduces gains of white-tailed doe fawns. Some of these effects on average daily gain in sheep are the result of increased gut fill and pelt weight. Increasing daily exposure to light increases feed intake when sheep or cattle are fed ad libitum. However, increased feed intake is not a prerequisite for the anabolic effects of long duration exposures to light because increased growth occurs in the animals given 16 h light:8 h dark (16L:SD) even when feed intake is restricted. The anabolic effects of increased duration photoperiods in sheep are independent of the gonads, whereas in cattle they are dependent on the gonads. Consistent increases in average daily gains of cattle in response to longer duration photoperiods have not always been achieved. The lack of consistency may be associated with sexual maturity or rate of fattening of the animal. For example, the stimulatory effects of 16L:8D photoperiods on live weight gain are not readily manifested in immature prepubertal heifers, but occur primarily during the peripubertal period. Short days are conducive to deposition of fat, which may account for the stimulatory effects of short days on live weight

gain of white-tailed doe fawns and excessively fattened Holsteins. In contrast, long duration photoperiods stimulate protein accretion in cattle. The hormonal signals that mediate the anabolic effects of increasing exposure to light are not associated with change in insulin, thyroxine or growth hormone concentrations in the blood. Glucocorticoid concentrations in serum decrease with longer duration photoperiods which is consistent with an anabolic effect. Increasing daily light exposure to 16 h/d hastens the increase in concentrations of progesterone and testosterone in sera of peripubertal heifers and prepubertal bulls, respectively. Thus, change in secretion of reproductive hormones in the peripubertal period of cattle may be associated with the anabolic effects previously described and is consistent with gonad dependency. Prolactin concentrations in serum increase in sheep and cattle as duration of light exposure increases. But, there is no irrefutable proof that prolactin is responsible for the increased rates of gain or change in carcass composition associated with longer exposures to light. Thus, the hormonal mechanisms involved in photoperiod-induced alterations in growth remain to be established. (Key Words: Photoperiod, Body Weight Gain, Body Composition, Nutrient Intake, Hormones.)
Introduction

At least two climatic variables affect rates of t Paper presented at the symposium on "Red body growth and reproduction in mammals. Meat Production and Processing Systems for the 21st For example, ambient temperatures beyond the Century" at the 75th Annu. Meet. of the Amer. Soc. Anita. Sci., Washington State Univ., Pullman, July thermoneutral zones of animals reduce growth 29, 1983. rates and reproductive efficiency. In addition, 2Michigan Agr. Exp. Sta. Journal Article No. daily photoperiod has long been known to 11035. This research was supported in part by USDA affect reproductive efficiency of many species. Grants 901-15-2 and 70-59-2261-1-2-O72-1 and USPHS Grant HD 17738. As a result, there has been a sharp increase over SOn leave from Lennoxville Res. Sta., Agriculture the last several decades in housing domestic Canada, Lennoxville, Quebec. food-producing animals in close confinement, 4Dept. of Anim. Sci. thereby permitting more regulation of their Received December 12, 1983. environments and enhancing their productivity. Accepted May 11, 1984. 1610 JOURNAL OF ANIMAL SCIENCE, Vol. 59, No. 6, 1984
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PHOTOPERIOD, BODY GAIN AND COMPOSITION The focus of this article will be on photoperiodic regulation of growth. Currently, most commercial applications of lighting schemes for domestic farm mammals are designed to provide light throughout the night. Recent studies suggest, however, that continuous lighting does not stimulate nutrient intake or growth rates. In this paper we will describe results of experiments that show controlled daily light (L):dark (D) cycles affect body growth, carcass composition, nutrient intake and hormone secretion in sheep, deer and cattle. There is no good evidence that photoperiod affects growth of pigs (Mahone et al., 1979; Ntunde et al., 1979; Berger et al., 1980; Hoagland and Diekman, 1982); and to our knowledge, there are no published data for the horse. Therefore, these classes of livestock are excluded from this discussion.
Body Growth and Carcass Composition

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Forbes et al. (1975, 19795) first reported an increase in live weight gain of castrated male or intact ewe lambs exposed to 16L:8D photoperiods as compared with animals exposed to 8L: 16D. One-half the lambs were fed a concentrate diet ad libitum, whereas the other one-half were restricted to gain 70 g/kg body weighC 'Ts" head - i ' d -1. In comparison with 8L:16D, 16L:SD stimulated live weight gain of lambs fed ad libitum (230 vs 204 g/d) and restricted (154 vs 123 g/d) diets (figure 1). In pair-fed animals, live weight gain was significantly greater under 16 h daylengths, but most of these differences were ascribed to increased gut fill and pelt weight (Forbes et al., 1979b). in addition to live weight gains, 16L:8D increased carcass weight of sheep (Forbes et al., 1975, 1979b; Schanbacher and Crouse, 1980, 1981). Based on physical separation of the 11 to 13th rib section, it was concluded that 16L:8D induced increments in carcass weight gain were of normal composition, not an increase in fat percentage (Forbes et al., 1975 ; 1979b). In a few experiments, the carcasses of sheep exposed to 16L:8D were leaner than those of sheep given 8L:16D (Forbes et al., 1979b; Schanbacher et al., 1982). In other experiments, photoperiod did not affect overall measures of backfat thickness, kidney and pelvic fat or quality and yield grade in sheep (Schanbacher and Crouse, 1980, 1981). In addition, Forbes et aI. (1979b) reported that

16L:8D exposures increased muscle mass in the carcass and did not change fat percentage in the 11 to 13th rib section or in the fat depots. Anabolic effects of 16L:8D on live weight gain, carcass weight and composition were observed in lambs regardless of ambient temperature (5, 18 and 31 C) during the rearing period (Schanbacher et al., 1982). However, chemical analysis of the carcasses of sheep have not yet been reported. Thus, definitive conclusions concerning photoperiodic control of carcass composition in sheep remain to be established. Similar to sheep, deer become reproductively active during photoperiods of short duration. In contrast to sheep, however, short duration photoperiods were stimulatory to increases in live weight gain (Budde, 1983; Abbott et al., 1984). For example, white-tailed doe fawns exposed to 8L:16D for 2 or 3 mo exihibited larger average daily gains than fawns given 16L:8D (table 1). Weights of carcass, skeletal muscle !including intramuscular fat) and viscera of short-day exposed fawns were considerably heavier than those of fawns exposed to long days. These increases in weight in response to short photoperiods were attributed primarily to increased deposition of fat. Coincident with the increased percentage of fat, there was a decreased percentage of protein in the carcasses. In cattle, live weight gain increased 11 to 17% in heifers exposed to 16L:8D over that of heifers exposed to natural duration photoperiods of 9 to 12 h daily (Peters et ah, 1978) or 8L:16D (Peters et al., 1980; Petitclerc et al., 1983a). Even when feed intake was restricted and identical (diet designed to permit gains of .7 kg/d) for groups of heifers exposed to 8L:16D or 16L:8D, increased rates of weight gain persisted in animals given the longer exposure to light (figure 2; Petitclerc et al., 1983a). In fact, the percentage increase in weight gain was greater in the restricted-fed heifers than in heifers fed a diet ad libitum to provide gains of 1 kg/d, which agrees with data for sheep (figure 1; Forbes et al., 1979b). Petitclerc et al. (1984) was first to use chemical analyses to address the effects of photoperiod on carcass composition of cattle. A photoperiod of 16L:8D enhanced gain in percentage of protein (11.0%) in the 9 to 11th rib section of heifers fed ad libitum a diet formulated to provide gains of 1 kg/d, but not in heifers fed a diet designed to provide gains of .7 kg/d (figure 3). Percentages of water and

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TUCKER ET AL.
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Figure 1. Live weights of lambs exposed to 16 or 8 h of light/d and fed ad libitum (AL) or restricted (R) quantities of concentrates. From Forbes et al. (1979b).

fat in the carcasses were not affected by photoperiod treatments. During winter, continuous lighting was less effective in stimulating b o d y growth than natural short-day photoperiods in sheep (Hulet et al., 1968). In cattle, continuous lighting was less effective than 16L:SD (figure 4; Peters et al., 1978). Therefore, a period of dark appears necessary for maximal stimulation of growth. However, light does not need to be present as a continuous block of 16 h to stimulate growth. Long duration photoperiods of 16L:8D can be mimicked b y coupling short durations of light with a brief "flash" of light given at a precise time during the dark period. For example, lambs exposed to a photoperiod of 7 L : 9 D : I L : 7D gained weight more rapidly than lambs

exposed to 8L:16D (table 2; Schanbacher and Crouse, 1981; Brinklow and Forbes, 1982). Similarly, in cattle, heifers exposed to 6L:SD: 2L:8D tended to grow faster (3%) than heifers exposed to 8L:16D (S. A. Zinn and H. A. Tucker, unpublished data). These reports support the hypothesis of Bunning (1960) for a diurnal rhythm of photosensitivity to light. Physiological responses such as stimulation of growth occur when light coincides with a specific phase of an endogenous rhythm of photosensitivity. I n cattle and sheep, stimulation of growth is expressed when light is present 14 to 17 h after dawn (or 8 to 10 h after dusk). When light is not given during this photosensitive phase, such as in the experiments of Hackett and Hillers (1979), there is no stimulation of growth. The role of the gonads on photoperiodinduced growth differs between sheep and cattle. In sheep, Schanbacher and Crouse (1980) observed that long-duration photoperiodic stimulation of growth was similar in 10-wk-old castrated and intact ram lambs (table 3). Thus, the b o d y growth response to photoperiod in sheep is independent of the gonads. On the other hand, exposure of young immature male calves (10 to 14 d old) or 15-mo-old steers to 16L:8D for 84 to 100 d did not stimulate b o d y or carcass growth over that obtained with 8L:16D or natural short-day winter photoperiods in Ireland (Roche and Boland, 1980). Similarly, we did not observe any differences in average daily gain of finishing steers exposed to

TABLE 1. SELECTED GROWTH CHARACTERISTICS OF WHITE-TAILED DOE FAWNS EXPOSED TO 16L.-8D OR 8L: 16D PHOTOPERIODS FOR 14 WKa Trait Average daily gain, g Carcass weight, kg Skeletal muscle Weight, kg Crude protein, % Fat, % Viscera Weight, kg Crude protein, % Fat, % Muscle plus viscera Protein, g Fat, g aFrom Abbott et al. (1984.). 16L:8D 161 18.7 10.6 80 12 3.6 64 29 2,180 621 8L:16D 113 15.4 13.5 61 34 4.4 40 54 3,226 2,269

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PHOTOPERIOD, BODY GAIN AND COMPOSITION

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Figure 2. Live weights of heifers exposed to 16 or 8 h of light/d and fed ad libitum or restricted diets. Average daily gains are s h o w n to the right for each treatment. From Petitclerc et al. (1983a).

16L:8D or natural short-day photoperiods during winter and spring in Michigan (H. A. Tucker, D. Petitclerc and L. T. Chapin, unpublished data). However, 16L:8D increased average daffy gain 9.8% in intact Holstein bulls, whereas there was no significant effect in Holstein steers (figure 5). Thus, photoperiodinduced growth of cattle appears to be a gonad-dependent phenomenon. In addition to the gonads, initial body weight of the animals may play a role in the photoperiodic response. During winter-spring and autumn-spring, two experiments were conducted to investigate the effect of 16L:8D in comparison with natural photoperiod on body weight gains of perpubertal Holstein heifers (n=68 and 64, respectively). In order to reduce competition for feed availability between small and large animals, heifers were divided into two groups according to body weight at the beginning of both trials. In both experiments, 16L:8D stimulated body growth of small size heifers by 9.2 and 9.1%, respectively, but did not influence growth rate of large size heifers (figure 6). Most of the large size heifers reached puberty relatively soon after beginning the experiment, whereas the smaller heifers reached puberty towards the end of the experiment. Thus, increments in body growth in response to 16L:8D photoperiods were best achieved when the prepubertal heifers were

approximately 110 kg at the beginning of the experiment and photoperiod treatments continued through puberty. Collectively, the cattle studies led us to postulate that the degree of maturity of the gonads could play a significant role in the growth response to photoperiod. In order to investigate this question further, Zinn et al. (1983b) subjected prepubertal (84 kg average body weight at beginning of experiment) heifers to either 8 or 16 h of light daily from October to February. In heifers that remained prepubertal, 16L:8D nonsignificantly stimulated average daily gain 4% over that of similar heifers given 8L:16D. However, the difference widened as the experiment progressed. Perhaps with longer exposure the overall differences would have become greater if the animals had been allowed to proceed through puberty. In contrast to the response of prepubertal heifers, in heifers postpubertal at the beginning of the experiment a photoperiod of 8L:16D stimulated average daffy live weight gains 13% over that of the postpubertal heifers exposed to 16L:8D. However, a high energy diet was fed ad libitum and designed to provide an average daffy gain of 1 kg in the prepuberal heifers. The same diet was fed ad libitum to postpubertal

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Figure 3. Percentage of protein in 9 to l l t h rib section of heifers fed a low or high plane of nutrition and exposed to 8 or 16 h of light/d. Heifers were killed at an average body weight of 345 kg after 181 to 236 d on experiment. F r o m Petitclerc et al. (1984).

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TUCKER ET AL.

TABLE 2. GROWTII AND FEED INTAKE OF RAM LAMBS EXPOSED TO VARIOUS PHOTOPERIODSa Photoperiod 8L:16D 16L:8D 7L:9D:IL:7D aFrom Schanbacher and Crouse (1981). Avg daily gain, g/d 345 417 442 Feed intake, kg 125 141 138

heifers and this led to growth rates greater than 1.1 kg/d. These postpubertal Holstein heifers gained excessive amounts of fat. Chemical analysis of the 9 to l l t h rib section revealed that 8L: 16D stimulated fat deposition 8% more than that in ribs of postpubertal heifers given 16L:8D. In contrast, protein in ribs of postpubertal heifers given 16L:8D averaged 15%, whereas heifers exposed to 8L:16D averaged 14%. We postulate that short duration photoperiods favored fat deposition in heifers, and this accounted for their more rapid gain. In contrast, heifers given 16L:8D diverted greater amounts of their nutrients to the energetically less efficient process of protein accretion. Thus, our current working hypothesis is that short days are stimulatory to fat accretion in cattle and especially deer, whereas long days stimulate protein accretion.
Feed Intake

Continuous night lighting of cattle has been practiced commercially for many years, presumably to stimulate eating activity. However, evidence to support this assumption is not great. For example, Holstein heifers 3 to 5 mo of age exposed to 16L:8D for 16 wk consumed 6.9% more dry matter/d than heifers exposed to continuous lighting (24L:0D) or to less than 11.6 h of light daily (figure 7; Peters et al.,

1980). In addition, lactating Holstein cows given 18L:6D had as many eating events as cows given continuous illumination, and milk production did not differ in the two groups (Tanida et al., 1984). Because continuous night lighting does not stimulate feed intake over that obtained with 16L:8D, there is little reason to employ continuous lighting to increase eating activity. Hence, costs of lighting could be reduced. There may be, of course, reasons other than feed intake to employ continuous night lighting. When fed moderate energy density diets, sheep and heifers exposed to 16L:8D grew faster and consumed more dry matter than similar animals reared under 8L:16D photoperiods (Forbes et al., 1975; Schanbacher and Crouse, 1980; Peters et al., 1980; Petitclerc et al., 1983a). Nonetheless, 16L:8D photoperiods induced greater efficiency in terms of feed to gain ratio. Sheep given photoperiods of 7L:9D: 1L:7D consumed less feed per unit of body weight gain than control animals given 8 h of light as a continuous block each day (table 2; Schanbacher and Crouse, 1981).

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TABLE 3. AVERAGE DAILY GAINS OF RAM AND WETHER LAMBS EXPOSED TO 16 OR 8 H OF LIGHT DAILYa Treatment Ram (16L:8D) Ram (8L:16D) Wether (16L:SD) Wether (8L: 16D) Avg daily gain, g/d 410 340 345 300

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aFrom Schanbacher and Crouse (1980).

Figure 4. Live weights of Holstein heifers exposed to 16L:8D, 24L:0D or natural photoperiods between November and March. From Peters et al. (1980).

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PHOTOPERIOD, BODY GAIN AND COMPOSITION

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Figure 5. Live weights of Holstein bulls and steers exposed to 16L:8D or natural photoperiod (n = 13/treatment).

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Figure 6. Live weights of Holstein heifers exposed to 16L:BD or natural photoperiod. In each experiment, animals were blocked into groups of eight heifers each according to initial body weight. Within blocks heifers were assigned randomly to treatment. Heifers were subsequently housed in four separate pens, two pens for largest heifers, and two pens for the smallest.

24L:OD Natural D~v~en~h Nov 11 to March 10

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Figure 7. Dry matter intake of Holstein heifers exposed to 16L:8D, 24L:OD or natural photoperiods between November and March. From Peters et al. (1980).

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TUCKER ET AL. period and hormone secretion on body growth, carcass composition and utilization of nutrients have not been established. Nevertheless, a reasonable body of data has accumulated to permit formulation of working hypotheses as to which hormones may or may not be involved. Several hormones normally considered to be involved in growth or diversion of nutrients among various organs are probably not involved in photoperiod-induced changes in growth. These would include growth hormone, insulin and thyroxine (table 4). To our knowledge, photoperiod does not affect secretion of these hormones in the red-meat animals described in this article. For example, in cattle, mean serum concentrations, thyrotropin-induced releases, clearance rates and secretion rates of growth hormone were similar after several weeks exposure to 8 or 16 h of light/d (Leining et al., 1980; S. A. Zinn, L. T. Chapin and H. A. Tucker, unpublished data). Similarly, mean concentrations of thyrotropin, thyroxine and insulin in serum were unresponsive to changing photoperiods in cattle (Leining et al., 1980; K. B. Leining and H. A. Tucker, unpublished data). Similar to findings in cattle, growth hormone, insulin and thyroxine concentrations in serum of sheep were unaffected by photoperiod (Forbes et al., 1979a). In deer, numbers of growth hormone secreting cells did not change during the seasons (Schulte et al., 1980b). Leining et al. (1980) observed in prepubertal bulls that basal concentrations of glucocorticoids in serum decreased as duration of light was increased from 8 to 16 h/d (table 4). Conversely, glucocorticoids increased 118% when daily light was decreased from 16 to 8 h. Because glucocorticoids are generally catabolic and negatively associated with rates of gain (Purchaset al., 1971), it may be hypothesized that decreased secretion of these hormones are

secondary to effects on body growth. Moreover, as described in the previous section, restriction of feed intake did not preclude the growth promoting effects of supplemental lighting in sheep or cattle (Forbes et al., 1979b; Petitclerc et al., 1983a). These findings suggest that stimulatory effects of photoperiod on growth are not mediated primarily through increased voluntary feed intake. Light-dark cycles affect eating patterns. Sheep and heifers exposed t o 16L:8D had more eating events and showed more of these events in lighted hours than animals treated with 8L:16D (Schanbacher and Crouse, 1981; Zinn et al., 1983a). Heifers on 8L:16D increased their eating activity 1 to 2 h before lights came on, whereas heifers on 16L:SD did not reinitiate eating activity until lights were actually on. Offering fresh feed also stimulated eating activity. Thus, lights on and presentation of fresh feed increased eating events. However, eating pattern has not been directly linked to rates of body weight gain, except that feeding once per day within 1 h before lights out reduced subsequent body gain in comparison with feeding 1 h after dawn or feeding in the middle of the day (Zinn et al., 1983a).
Postulated Hormonal Mechanisms

In seasonal breeding mammals, photoperiod markedly affects secretion of the classical hormones of reproduction: luteinizing hormone, follicle-stimulating hormone, estradiol, progesterone and testosterone. These responses have been summarized elsewhere (Karsh and Foster, 1981; Tucker and Ringer, 1982) and will only be described in this article as they may pertain to growth, not reproduction. Emphasis will be placed on those hormones closely associated with metabolism and growth. Cause and effect relationships between photo-

TABLE 4. CONCENTRATIONS (NG/ML) OF SELECTED HORMONES IN HEIFERS EXPOSED TO 16 OR 8 H OF LIGHT DAILY FOR 6 WKa Horrnone Growth hormone Thyrotropin Thyroxine Insulin Glucocorticoids 16L: 8D 9.5 3.5 60 2.3 1.5 8L: 16D 8.8 3.2 56 2.3 2.8 Level of probability P>.05 P>.05 P>.05 P>.05 P<.05

aFrom Leining et al. (1980) and K. B. Leining and H. A. Tucker (unpublished data).

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PHOTOPERIOD, BODY GAIN AND COMPOSITION

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involved in the photoperiodic-induced changes in growth and carcass composition. Additional research in which blood samples are collected throughout the day and night will be needed, however, to eliminate the possibility that diurnal variation in concentrations of glucocorticoids accounts for the apparent differences in concentration of glucocorticoids in prepubertal bulls given 8 or 16 h of light/d. Testosterone is probably not involved in photoperiod-induced changes in growth of rams, but it remains a possible mediator in bulls. In sheep, long-duration photoperiods stimulated body growth but inhibited secretion of testosterone (Lincoln and Peer, 1977; Schanbacher and Ford, 1979). Moreover, 16L:8D photoperiods stimulated castrated rams to grow faster than castrates given 8L:16D (Forbes et al., 1975). Thus, the growth response of sheep to photoperiod is independent of gonadal steroids. Photoperiodic-induced increments in growth of cattle may be associated with changes in secretion of hormones at puberty. Indeed, puberty in heifers and bulls (based on increased concentrations of progesterone and testosterone in serum, respectively) occurred earlier in animals exposed to 16L:8D than in animals given 8L:16D (Peters and Tucker, 1978; Stanisiewski et al., 1982; Petitclerc et al., 1983a), and coincided with photoperiod-induced increases in growth. In contrast to sheep, castrate bulls were unresponsive in terms of body growth to change in photoperiod. Thus, gonadal hormones are likely to be involved in photoperiod-induced growth of cattle. Of the metabolic hormones studied, prolactin is the most responsive to changes in photoperiod in ruminants. Initial studies in cattle showed that gradually increasing daily exposure from 8 to 16 h of light increased prolactin concentrations in serum fourfold (Bourne and Tucker, 1975). Conversely, reducing duration of daily light from 16 to 8 h decreased concentrations of prolactin 86%. Furthermore, abrupt increases in light exposure from a continuous block of 8 h of light/d to 16 continuous h of light/d increased basal concentrations of serum prolactin 2- to 10-fold in cattle [Leining et al., 1979; Stanisiewski et al., Petitclerc et al., 1983b (figure 8)]. However, light does not necessarily have to be supplied as a continuous daily block to stimulate secretion of prolactin. For example, a photoperiod of 7 L : 9 D : I L : 7 D was as effective as 16L:8D in stimulating basal

secretion of prolactin in rams and ewes (Ravault and Ortavant, 1977; Thimonier et al., 1978). Insertion of a 1-h pulse at other times during the dark period was ineffective in stimulating secretion of prolactin. Similarly in cattle, 6L:8D:2L:8D increased basal concentrations of prolactin in serum to the same degree as 16L:SD (figure 8; Petitclerc et al., 1983b). A photoperiod o f 6 L : 14D:2L:2D was much less effective than 6L:8D:2L:SD in stimulating secretion of prolactin. Thus, the data support the hypothesis that sheep and cattle possess a photosensitive rhythm for secretion of prolactin, which also coincides with the previously described daily photosensitive period for stimulation of growth. Initial studies showed that when light periods were gradually increased from 8 to 24 h (continuous light) daily, concentrations of prolactin in serum increased 2.3-fold, but within a week after attaining 24L:0D, prolactin concentrations decreased 50% (Leining et al., 1979). More recent studies have shown that when photoperiods are abruptly switched from 8L:16D to 24L:0D prolactin increases approximately 250% (E. P. Stanisiewski, L. T. Chapin and H. A. Tucker, unpublished data). This increase was comparable with that observed when cattle were switched from 8L:16D to 16L:8D. However, prolactin concentrations in serum attained their maximum about 3 wk sooner in animals given 16L:8D than when switched to continuous light. In these experiments, increased prolactin concentrations were

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Figure 8. Prolactin concentrations in serum of propubertal bulls exposed for 6 wk to 8L:16D (open symbols) then switched to 16L:8D or 6L:8D.-2L:8D for 6 additional wk (closed symbols). Onset of lights during the primary period of lighting was 0700 h. From Petitclerc et al. (1983b).

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TUCKER ET AL.

maintained with either 16L:8D or 24L:0D. De- beginning of each dark period each day (Ravault spite maintaining a similar level of prolactin, a and Ortavant, 1977; Lincoln, 1979). The photoperiod of 24L:0D did not stimulate physiological significance of this dark-induced increased rates of live weight gain as did 16L:SD pulse of prolactin in sheep remains to be (Peters et al., 1980). determined. Sixteen hours of light/d from red, blue or There is some circumstantial evidence that Vita-Lite| fluorescent, incandescent, mercury prolactin may not be involved in growth. For vapor or high pressure sodium lamps were as example, exposure to continuous lighting, effective as cool-white fluorescent light in stimulated secretion of prolactin (E. P. stimulating secretion of prolactin (Leining et Stanisiewski, L. T. Chapin and H. A. Tucker, al., 1979; Stanisiewski et al., 1984). Whether all unpublished data), but did not affect body of these lamp types will stimulate growth rates growth rates (Hulet et al., 1968; Peters et al., has not been determined, although we have 1980). But whether or not continuous light some very preliminary data that Vita-Lite| maintains increased concentrations of prolactin may be superior to cool white fluorescent in the period required to observe increased lamps in stimulating milk production as well as growth rates is unknown. In addition, suppresrates of gain (E. P. Stanisiewski, S. A. Zinn sion of secretion of prolactin with ergot alkaand H. A. Tucker, unpublished data). loids failed to affect body growth of sheep As summarized by Forbes (1982), intensities (Ravault et al., 1977). However, the effects of of artificial light greater than 100 lx may be ergot alkaloids on growth of animals exposed to essential to achieve enhanced rates of growth. long or short day photoperiods has not been Light intensities of approximately 200 to 600 reported. As ambient temperatures decrease, lx applied for 16 h each day were also effec- the ability of 16 h photoperiods to increase tive in increasing serum concentrations of prolactin concentrations in serum was reduced prolactin (Peters and Tucker, 1978; Leining et (Peters and Tucker, 1978), yet the increased al., 1979). Intensities outside this range may be rates of gain persisted in cold environments effective, but responses have not been reported. (Peters et al., 1978; Peters et al., 1980). These The ability of 16 h of light to stimulate data should be interpreted cautiously, however, secretion of prolactin above that found in because neither temperature nor prolactin were animals exposed to 8 h of light/d also occurs in measured continuously, and unrecorded sporasheep (Forbes et al., 1975; Ravault and Orta- dic increases in temperature could have allowed vant, 1977). In white-tailed deer, prolactin photoperiod-induced increments in prolactin concentrations were greatest in summer and secretion to be expressed. It is possible that lowest in winter (Schulte et al., 1980a), but sporadic increases in secretion of prolactin may whether these responses are associated with be sufficient to accelerate anabolic processes. photoperiod, ambient temperature or a combi- There is precedence for this possibility. In dairy nation has not been established. In contrast to cows, the temporary (2 to 3 d) increase in the data in ruminants, photoperiod had no secretion of prolactin during the periparturient effect on concentrations of prolactin in pigs period was sufficient to cause lactogenesis (Hoagland et al., 1981). This agrees with the (Akers et al., 1981), a process that occurs over fact that photoperiod does not stimulate several weeks. Yet, during established lactation, growth rates of pigs. continuous high secretion of prolactin was not After an abrupt change in photoperiod, the required for maintenance of milk production in first detectable alterations in concentrations of cows (Smith et al., 1974). prolactin in serum usually did not occur for Thus, prolactin may not yet be ruled out as approximately 1 wk in cattle or sheep and a mediator of photoperiodic-induced changes in maximal concentrations were not achieved for growth of ruminants. Indeed, there is some several weeks (Lincoln et al., 1978; Leining et evidence that prolactin may be involved in al., 1979). After maximal or minimal concen- anabolism. For example, prolactin administratrations of prolactin were reached when exposed tion induced anabolic responses in a variety of for several weeks to 16L:8D or 8L:16D, there species (Nicoll, 1980). Infusion of prolactin was no obvious diurnal pattern in secretion of increases N retention in sheep maintained in prolactin in cattle (figure 8; Petitclerc et al., continuous darkness (Brinklow and Forbes, 1983b). Sheep, in sharp contrast, markedly 1983). Conversely, immunizing sheep against increased their secretion of prolactin at the prolactin suppressed body growth rates (Ohlson

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PHOTOPERIOD, BODY GAIN AND COMPOSITION et al., 1981). Moreover, p i n e a l e c t o m y b l o c k e d p h o t o p e r i o d - i n d u c e d changes in p r o l a c t i n as well as p h o t o p e r i o d - i n d u c e d i n c r e m e n t s in g r o w t h rate o f sheep ( B r o w n and F o r b e s , 1980). Obviously, a d d i t i o n a l research will be n e e d e d to establish t h e h o r m o n a l c o n t r o l o f p h o t o p e r i o d - i n d u c e d changes in rates o f gain and carcass c o m p o s i t i o n .

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