Posthatch Development of the Gastrointestinal Tract and other Organs of Chickens.

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1 Posthatch Development of the Gastrointestinal Tract and other Organs of Chickens. Zafrira Nitsan Agricultural Research Organization, The Volcani Center, P.O.Box 6 Bet Dagan, Israel Age of market weight of broilers has been continuously reduced by approximately one day per generation during the past years due to an average increase of 50 g/year of body weight at 42 days of age (Gyles, 1989). Although non-genetic factors such as improved feeding and management contributed to this growth rate increase, genotypic selection for growth seems to be the major cause (Chambers et al., 1981; Marks, 1979). This trend emphasizes the importance of the early growing period in today's broiler; increasing the proportion of the first week after hatch from 8% to 16% of the broiler's life span in the last 30 years. The decreasing growing period allows less time (which is reduced with each new generation) for compensation for any growth lag period, therefore any factor limiting maximal early growth delays marketing and reduces efficiency of production. The importance of growth rate during the first week of life is further emphasized by the significant high correlation between body weight at 42 and 6 days of age, although there is no correlation of 42 day weight with body weight at hatching. Growth curves of broiler and Leghorn chicks (Fig. i) and of different genetic stocks (Katanbaf et al., 1988) show a growth lag of several days after hatch. Fig. i. BODY WEIGHT AND FOOD INTAKE OF BROILER AND LEGHORN CHICKS DURING 15 DAYS AFTER HATCH (0 -broilers; 0 -Leghorns)... BODY.WEIC_.M-IT F:_.LATIVE C_9::{OWTH FOOD INTAKE RELATIVE FOO0 INT/4< E /t a_.. 1o "; O0 " O ".. 3 az S_.3 4 G I_ IS IZ *S G 3 6 D I $ AGE (days) According to: Nir, Nitsan, Mahagma, (in press). The relative growth of the two breeds was very low immediately after hatch and gradually increased to a peak of approximately 14% 32

2 for Leghorns and 20%/day for broilers 4 to 5 days after hatch. Age and height of these peaks may vary with stock, food composition and management. Comparing the relative growth of boiler chicks with faster growing species of birds, one can see differences not only in the magnitude of growth rate, but also in growth patterns (Fig. 2). While broilers reach gradually the peak of their relative growth, quail and squabs have the highest relative growth on the first day after hatch. Later on, while broilers maintain a relative daily weight gain of > 10% for at least 20 days, that of the two other species reduces markedly to values much below 10%/day and they reach mature weight between 21 to 28 days of age. Fig. 2. RELATIVE GROWTH OF SQUABS, QUAIL AND BROILER CHICKS _oilcr ---- squab... _ail O0 \ \ \ \ \ "_ 40 O 30 \ \, \ \ \ lo ': age (days) The close relationship between food intake and body-weight gain existing in young animals raises a hypothetical question: which factor is the mandatory one for expressing growth potential of different breeds or lines of birds - appetite or food utilization? Or, in other words, which is the limiting factor: appetite, which limits the amount of metabolites available to the tissues for growth, or the capacity of the digestive tract to digest and absorb efficiently large amounts of nutrients and the anabolic capacity of the tissues. Behavioral factors, associated with learning of eating and drinking, may also be involved in determining growth rate after hatch. The role of the vitelline residue in supplementing nutrients after hatch is limited and brief (Murakami et al., 1988; Nitsan et al., 1991). Its contribution is approximately 50 and 40% of the total energy and protein supplies (from food and yolk) on the first day after hatching and negligible by day 4 being only 2 and 6% of energy and protein, respectively. Therefore, a transition of the digestive system from embryonic absorption of yolk to ingestion and digestion of feed is essential for maximization of early growth. At the same time metabolic transition occurs; while in embryonic stage most of the energy is supplied by fat (from the yolk), the main energy source of the 33

3 posthatched chick is carbohydrates (from grains). The main theories considered to be closely associated with food intake regulation are: Glucostatic theory, which does not hold for birds as for mammals. Although blood glucose level is 1.5 to 2 times higher in birds than in mammals, it seems that it has little effect on food regulation. Increasing blood glucose level by injection of glucose or glucagon hardly affects food intake and insulin suppresses intake, a response contrary to that seen in most mammals (Smith & Baranowski-Kish, 1979). Lipostatic theory seemed to apply better in birds, in which the ability of the adipose tissues to incorporate surplus fatty acids, resulting from excessive food absorbed, may be a limiting factor in food intake (Lepkovsky, 1973). Moreover, it was suggested that adipose tissue was also involved in the disposition of chickens to be overfed. It was shown that after a period of force feeding, which enhanced fat accumulation, food intake was low. It gradually increased during a period of ii to 13 days when the chicks resumed food intake at the level of their ad libitum-fed counterparts. During this period the chicks lost weight and probably the extra fat accumulated while they were force-fed (Nir et al 1978) i.e fat stores may affect food intake. This theory is consistent with observations in crammed geese in which, after reaching the set point of fat content in adipose tissues and blood, there is an interruption in the passage of food in the digestive tract, probably as the last defensive mechanism against lethal obesity. Selection for body weight or for adiposity may change the lipostatic set point. Food intake of ad libitum-fed broiler chicks is much greater than that of Leghorn chicks, although the former have heavier adipose tissues. Moreover, chicks selected for high or low abdominal fat, in which the differences in adiposity is much greater than between egg and meat-type chicks, consume similar amounts of food (Keren-Zvi et al., 1990). The large fat stores in the liver and residual yolk (approximately 20% in each, Nitsan et al., 1991) and the high blood lipid levels (i0 mg/ml) in chicks at hatch, may be one of the reasons for the lag time between hatching and the start of eating. Most of these stores are depleted during the first three days after hatch. Eighty percent of the yolk's fat is used on the first day (Nitsan et al., 1991). With the recognition that food intake is not dependent on a single mechanism but is influenced by many factors, of which metabolic effects on hunger or satiety centers are probably the main long-term regulators, the digestive tract was suggested to be involved in short-term regulation of food intake. Distension of the gastro-intestinal tract, crop in particular, is believed to affect food intake. The presence of food in stomach and/or other parts of the intestinal tract, gives rise to satiety signals which negatively affect food intake. In order to meet energy needs, birds given diets diluted with inert materials increase food intake markedly to meet energy requirements, up to a limit, probably due to a feedback mechanism that arises in the digestive tract (Hill, 1978). In accord, for a long time there was a consensus that when given high density diets, food intake reduces because chickens 34

4 consume food according to their energy needs. However, the modern broiler consumes excessive amounts of energy as the dietary energy increases, indicating that the digestive tract and not appetite is the limiting factor in food intake. In squabs, appetite cannot be a limiting factor for growth, because the parents force feed them with excessive amounts of food, overcoming all satiety signals that might originate in a markedly distent crop. Shortly after hatch the crop with it's contents reaches 30% of body weight. In order to utilize this excessive amount of food, squabs exhibit extremely rapid growth of the pancreas and the liver and have maximal synthesis of pancreatic digestive enzymes at hatching or very closely thereafter. Contrary in chicks, the level of digestive enzymes at hatching is low and after gradual increases they attain maximal values only approximately i0 days after hatch. Selection for growth in broilers is tightly followed by selection for increased ad libitum intake (Burkhart et al., 1983). llowever, the modern broiler reached a point that closely after hatch it uses all its digestive system potential to ingest and metabolize food. "Bottlenecks" or Constrains affecting food intake and growth may be emphasized by exposing the birds to feeding regimes, in which (i) food is not available at all times, and (2) the chicks have to consume high amounts in relatively short period, to accommodate for periods in which food is unavailable. Overfeeding by intubation is an extreme example of feeding regimes mentioned above. It bypasses the appetite barrier and may provide information on whether the digestive tract is the limiting factor for enhanced food intake and growth. Young chicks from egg and meat type breeds of chickens show marked differences in their response 'to overfeeding. Whereas in egg-type breeds the amount of food which could be overfed reached 70% above that consumed by their ad libitum-fed counterparts, in broilers the maximal excess of food that could be overfed was only 13% (Nir et al., 1978). In parallel, chicks from a common source, divergently selected for low (LW) or high (HW) body weight (Siegel, 1978), could be overfed up to 76 and 15% above ad libitum intake, respectively (Barbato et al., 1984) Fig. 3; BODY WEIGHT OF AD LIBITUM OR OVERFED BROILER vs. LEGHORN AND HIGH WEIGHT (HW) vs. LOW WEIGHT (LW) CHICKS. BODY WEIGHT OF AD LIBITUM AND UoD_' _,,01Ls, _Y_,C,,TOr ol_l,_c,ioi_,c,,,c,<_,s _d.b O. OV_,_,.'_D OVERFED HW AND LW CHICKS _a _ Ab MI.,..i< ).A>-".-----" T.- _. N'._ / _. _. _" Oa_ on exploit /.,we... m Bach*to *t al [r e_ al., 1918 AGg (DA_S) 35

5 The excess amount of food that can be overfed, may stand for the "Assimilation Potential Reserve" of the breed or line of chickens, which was found to be almost nil in breeds selected for fast growth. During force feeding periods, increases in most segments of the digestive tract, including pancreas and liver, were much more pronounced in the Leghorns than in broilers and in chicks selected for high weight than for low weight (Table i). It seems that the limited propensity of the digestive tract to increases by additional food (above ad libitum intake) in faster growing lines prohibit overfeeding. This suggests also different limiting factors for food intake in the different breeds or lines: while in broilers and HW line the development of the intestinal tract and not appetite could be limiting in newly hatched chicks, in Leghorns and LW line, appetite rather than the intestinal tract may limit food intake and growth. TABLE i. INCREASE OF BODY AND ORGAN WEIGHTS DURING OVERFEEDING OF CHICKS FROM DIFFERENT GENETIC STOCKS (LB-Leghorn; HB-broiler; LW-low weight; HW-high weight) LB' HB!' LW' ' HW" Body Crop Proventriculus Gizzard i.i i.i Duodenum Small intestine Pancreas Liver According to: 'Nir et ai.,1978; "Barbato et ai.,1984). While selecting for increased body weight, the increase of the digestive system, mainly the anterior parts, where most of the digestion and absorption takes place, was inferior to that of body weight. Dror et al., (1977) showed that the relative weight of duodenum, jejunum, pancreas and liver were smaller in broiler than in Leghorn chicks closely after hatch. Fifteen years later a similar comparison showed almost no differences in the relative weights of liver and small intestine, while the pancreas was smaller in broiler than in Leghorn chicks (Nir et al., 1993). Although the former consumed (relative to body weight) about twice as much food as the Leghorns during the first week of life (Fig.l), this excessive intake, reflected by higher amount of contents in the small intestine, was not compensated for by increased weight of the intestine and pancreas nor by enhanced synthesis of digestive enzymes; Levels of enzymes in the pancreas and small intestine were lower in broiler than in the Leghorn chicks. After hatch the weight of pancreas, small intestine and liver increase faster than does body weight (Katanbaf et al., 1988; Nitsan et al., 1991; Nir et al., 1993), until reaching peaks of their relative weights. The fast growth of these organs and the accompanying increase of digestive enzyme levels in the pancreas and small intestine, are concomitant to increased growth rate of 36

6 chicks after hatch. Only after these organs (supply organs) had reached the peaks of their relative weight, could the "demand organs" ( muscles, skin, feathers) expose a pronounced growth (Katanbaf et al., 1988). It was shown above that broiler chicks, with limited enzymes levels after hatch have to cope with higher amounts of chyme than do Leghorn chicks. It may be assumed that supplementation of the diet with digestive enzymes or increases in density with readily digestible ingredients for the first few days after hatch, may compensate for the limitations of the digestive tract until it reaches its maximal relative size and digestive potential. A behavioral approach aiming in engorging the chicks to go to the feeders shortly after hatch may also enhance early growth. It was found lately that providing newly hatched chicks with three-day old "guide" chicks accelerated the time that chicks perform eating activity (Mahagna et al., in press). Combining nutritional and behavioral aspects may accelerate the time to reach maximal growth (approaching the shape of growth curves of fast growing species - squabs and quail), and allow full expression of the genetic improvement. References Barbato, G.F., Siegel, P.B., Cherry, J.A., Nir, I Selection for body weight at eight weeks of age. 17. Overfeeding. Poultry Sci., 63, Burkhart, C.A., Cherry, J.A., Van Krey, H.P., Siegel, P.B Genetic selection for growth rate alters hypothalamic satiety mechanisms in chickens. Behav. Genet., 13, Chambers, J.R., Gavora, J.G., Fortin, A Genetic changes in meat-type chickens in the last 20 years. Canad. J. of Anim. Sci., 61, Dror, Y., Nir, I., Nitsan, Z The relative growth of internal organs in light and heavy breeds. Br. Poult. Sci., 18, Gyles, N.R Poultry, people and progress. Poult. Sci., 68,1-8. Hill, K.J Physical effects of food in the digestive tract in relation to food intake. In: Food Intake Regulation in Poultry. Edit. Boorman, K.N. and Freeman, B.M. Edinburgh, Br. Poult. Sci. Ltd. pp Hill, K.J., Dansky, L.M Studies of the energy requirements of chickens i. The effect of dietary energy level on growth and feed consumption. Poult. Sci., 33, Katanbaf, M.N., Dunnington, E.A., Siegel, P.B Allomorphic relationships from hatching to 56 days in parental lines and F1 crosses of chickens selected 27 generations for high or low body weight. Growth, Develop. & Aging, 52, Keren-Zvi, S., Nir, I., Nitsan, Z., Cahaner, A Effect of dietary concentration of fat and energy on fat deposition in broilers divergently selected for high or low abdominal adipose tissue. Br. Poult. Sci., 31, Lepkovsky, S Hypothalamic-adipose tissue interrelationships. Fedn. Proc. Fedn. Anim. Socs. Exp. Biol., 32, Mahagna, M., Nir, I., Nitsan, Z Influence of the presence of three day oldchickens on the behavior of meat and egg-type 37

7 posthatched counterparts. (In press). Marks, H.L Growth rate and feed intake of selected and non selected broilers. Growth, 43, Murakami,H., Akiba, Y., Horiguchi, M Energy and protein utilization in newly-hatched broiler chicks-studies on the early nutrition of poultry. Jpn. J. Zootech., Sci. 59, Nir, I., Nitsan, Z., Dror, Y., Shapira, N Influence of overfeeding on growth, obesity and intestinal tract in young chicks of light and heavy breeds. Br. J. Nutr. 39, Nir, I., Nitsan, Z., Mahagma, M Growth and development of the digestive organs and some enzymes in broiler and egg type chicks after hatching. (In press). Nitsan, Z., Ben-Avraham, G., Zoref, Z., Nir, I Growth and development of the digestive organs and some enzymes in broiler chicks after hatching. Br. Poult. Sci., 32, Siegel, P.B Response to 20 generations of selection for body weight in chickens. XVI World's Poult. Congr. 10, Smith, C.J.V., Baranowski-Kish, L.L Mechanisms of energy intake in poultry. In: Food Intake Regulation in Poultry. Edit. Boorman, K.N. and Freeman, B.M. Edinburgh, Br. Poult. Sci. Ltd. pp

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