Posthatching Water and Feed Deprivation Affect the Gastrointestinal Tract and Intestinal Mucosa Development of Broiler Chicks

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1 2003 Poultry Science Association, Inc. Posthatching Water and Deprivation Affect the Gastrointestinal Tract and Intestinal Mucosa Development of Broiler Chicks A. Maiorka, E. Santin, F. Dahlke, I. C. Boleli, R. L. Furlan, 1 and M. Macari Faculdade de Ciências Agrárias e Veterinárias UNESP, Departamento de Morfologia e Fisiologia Animal, Jaboticabal, São Paulo, Brazil Primary Audience: Broiler Production Managers, Nutritionists, Researchers SUMMARY We studied the effect of feed and water deprivation on gastrointestinal tract and intestinal mucosa development of chicks at 24, 48, and 72 h posthatching. The treatments were water and feed ad libitum, water ad libitum and no feed, no water but feed ad libitum, and no water and no feed. The relative weight of the yolk sac was not influenced by the treatments. However, at 48 and 72 h posthatching, the relative weight of the liver increased, and the gizzard + proventriculus weight decreased in birds receiving feed ad libitum. An increase in jejunum and ileum relative weights and lengths was observed when the birds were supplied with feed and water. The lack of water produced the same effect as the lack of feed, both causing a higher number of villi per area with reduction in villus size, when compared with feed and water ad libitum treatments. The results of this study revealed that feed and water are able to affect intestinal villus development after hatching, indicating that both feed and water must be supplied to the chicks immediately after hatching. Key words: chick, feed, gastrointestinal tract, mucosa development, water 2003 J. Appl. Poult. Res. 12: DESCRIPTION OF PROBLEM The poultry industry is highly concerned about the effect of the interval between hatching and housing on chick development. One of the major effects of fasting between hatching and housing is the decrease in body fluids causing dehydration and impairing growth. At hatching, the chick digestive system is anatomically immature [1], and its functional capacity is not totally developed [2, 3]. Thus, after hatching, the gastrointestinal tract undergoes morphological and physiological changes, including increased surface area for digestion and absorption. The morphological changes involve increases in intestinal length, villus height, and density and, consequently, in the number of enterocytes and goblet and enteroendocrine cells [4, 5, 6]. The physiological changes are related to increased production of 1 To whom correspondence should be addressed: rlfurlan@fcav.unesp.br.

2 484 JAPR: Field Report TABLE 1. Effects of deprivation of water, feed, or both after hatching on liver, gizzard + proventriculus, yolk sac, duodenum, jejunum, and ileum weight, expressed as percentages of body weight, and duodenum, jejunum, and ileum lengths (cm) in chicks 24 h posthatching A Weight (% BW) Length (cm) Gizzard + Treatment Liver proventriculus Yolk sac Duodenum Jejunum Ileum Duodenum Jejunum Ileum With 3.32 ± 0.36 a 7.18 ± ± ± 0.46 a 1.55 ± ± 0.36 a 9.46 ± ± 2.55 a ± 2.21 a Without 3.05 ± 0.24 b 6.89 ± ± ± 0.28 b 1.23 ± ± 0.11 b 8.54 ± ± 1.40 b ± 2.33 b Water With 3.10 ± ± ± ± ± ± 0.41 a 9.12 ± ± ± 2.44 Without 3.26 ± ± ± ± ± ± 0.11 b 8.87 ± ± ± 2.52 Source of variation NS NS NS NS Water NS NS NS NS NS NS NS NS water NS NS NS NS NS NS NS NS NS P a,b Means within a column for each variable with no common superscript differ significantly by Tukey s test (P < 0.05).

3 MAIORKA ET AL.: GASTROINTESTINAL TRACT DEVELOPMENT 485 TABLE 2. Effects of deprivation of water, feed, or both after hatching on liver, gizzard + proventriculus, yolk sac, duodenum, jejunum, and ileum weight, expressed as percentages of body weight, and duodenum, jejunum, and ileum lengths (cm) in chicks 48 h posthatching A Weight (% BW) Length (cm) Gizzard + Treatment Liver proventriculus Yolk sac Duodenum Jejunum Ileum Duodenum Jejunum Ileum With 3.83 ± 0.63 a 7.12 ± 0.58 b 4.31 ± ± ± ± ± 1.25 a ± 2.96 a ± 2.24 Without 3.41 ± 0.32 b 8.28 ± 0.84 a 3.89 ± ± ± ± ± 0.81 b ± 2.26 b ± 2.10 Water With 3.68 ± ± ± ± ± ± ± ± ± 2.83 Without 3.58 ± ± ± ± ± ± ± ± ± 1.36 Source of variation NS NS Water NS NS NS NS NS NS NS water NS NS NS NS NS NS P a,b Means within a column for each variable with no common superscript differ significantly by Tukey s test (P < 0.05).

4 486 JAPR: Field Report TABLE 3. Interactions between feed and water treatments on jejunum and ileum weights, expressed as percentages of body weight, and ileum length (cm) in chicks 48 h posthatching A Water With Without Jejunum (% BW) With 2.70 ± 0.59 ay 1.54 ± 0.21 az Without 1.74 ± 0.42 by 1.71 ± 0.40 ay Ileum (% BW) With 1.77 ± 0.38 ay 1.01 ± 0.12 az Without 1.07 ± 0.19 by 0.98 ± 0.05 ay Ileum (cm) With ± 0.83 ay ± 2.71 az Without ± 1.26 by ± 1.52 ay a,b Means within a column for each variable with no common superscript differ significantly (P < 0.05). y,z Means within a row for each variable with no common superscript differ significantly (P < 0.05). pancreatic and intestinal digestive enzymes [2, 7] and membrane transporters [8], which carry the nutrients inside and later on outside enterocytes. The presence of nutrients in the intestinal lumen is able to stimulate villus and crypt growth [9]. However, the morphology of the intestinal epithelium is also particularly affected by the absence of food. Yamauchi et al. [10] reported that villus height in the duodenum was significantly reduced in birds exposed to 24-h fasting. The cell turnover of the intestinal epithelium was also significantly reduced in birds that fasted [4, 6]. The purpose of the present study was to evaluate the effects of posthatching water and feed deprivation on the gastrointestinal tract and mucosal development of broiler chicks. MATERIAL AND METHODS Seventy-two Cobb-500 male chicks from 60-wk-old breeders were used. Immediately after hatching, chicks were housed in an environmentally controlled room where ambient temperature was maintained at thermoneutrality (33 C) with continuous light. Chicks were distributed in battery pens with 18 chicks each divided into 4 groups: water and ration ad libitum; water ad libitum and no ration; no water and ration ad libitum, and no water and no ration. The diet used in the experiment was formulated based on corn and soybean meal with 22% of crude protein and 2,900 kcal ME/ kg. At 24, 48, and 72 h after hatching, 6 birds per treatment were slaughtered by cervical dislocation, and liver, gizzard + proventriculus, yolk sac, and small intestine were collected. The organs and each intestinal segment (duodenum, jejunum and ileum) were measured (cm ± 0.01) and weighed (g ± 0.01). To assure that birds received feed and water, gizzard + proventriculus and intestinal segments were opened to remove feed residues, which were observed in all fed birds, and water consumption was monitored by visual observations. Weights of the internal organs and intestinal segments were calculated as percentages of chick body weight. Samples of approximately 2 cm in length were collected from each segment of the small intestine (duodenum: distal region of duodenal loop; jejunum: region immediately anterior to Meckel s diverticulum, and ileum: region immediately anterior to the opening of the ceca), for analysis under a scanning electron microscope. The intestinal contents were removed with saline solution buffered with 0.1 M phosphate, ph 7.4, and the tissue samples were fixed in 2% glutaraldehyde in phosphate buffer for 24 h at 4 C. Subsequently, the tissue was washed in phosphate buffer and postfixed for 2 h in 1% osmium tetroxide. Next, the material was washed again with the same buffered solution and dehydrated in increasing ethanol series (30, 50, 70, 90, and 100% for 15 min each). Samples were dried in a critical point drier with

5 MAIORKA ET AL.: GASTROINTESTINAL TRACT DEVELOPMENT 487 TABLE 4. Effects of deprivation of water, feed, or both after hatching on liver, gizzard + proventriculus, yolk sac, duodenum, jejunum, and ileum weight, expressed as percentages of body weight, and duodenum, jejunum, and ileum lengths (cm) in chicks 72 h posthatching Weight (% BW) Length (cm) Gizzard + Treatment Liver proventriculus Yolk sac Duodenum Jejunum Ileum Duodenum Jejunum Ileum With 4.22 ± 0.39 a 7.80 ± 0.31 b 1.98 ± ± 0.46 a 2.42 ± ± ± ± ± 4.32 Without 3.55 ± 0.40 b 8.46 ± 1.02 a 1.94 ± ± 0.22 b 1.69 ± ± ± ± ± 2.10 Water With 3.97 ± ± ± ± ± ± ± ± ± 4.59 Without 3.84 ± ± ± ± ± ± ± ± ± 1.82 Source of variation NS NS Water NS NS NS NS NS water NS NS NS NS P a,b Means within a column for each variable with no common superscript differ significantly by Tukey s test (P < 0.05).

6 488 JAPR: Field Report TABLE 5. Interactions between feed and water treatments on jejunum and ileum weights, expressed as percentages of body weight, and duodenum, jejunum, and ileum length (cm) in chicks 72 h posthatching A Water With Without Jejunum (% BW) With 2.94 ± 0.60 ay 1.78 ± 0.09 az Without 1.89 ± 0.24 by 1.60 ± 0.20 ay Ileum (% BW) With 2.19 ± 0.17 ay 0.99 ± 0.11 az Without 1.24 ± 0.26 by 0.95 ± 0.08 ay Duodenum (cm) With ± 1.03 ay 8.75 ± 1.13 az Without 8.50 ± 0.71 by 8.33 ± 1.03 ay Jejunum (cm) With ± 6.68 ay ± 2.46 az Without ± 2.33 by ± 1.17 ay Ileum (cm) With ± 5.43 ay ± 1.38 az Without ± 1.38 by ± 2.02 ay a,b Means within a column for each variable with no common superscript differ significantly (P < 0.05). y,z Means within a row for each variable with no common superscript differ significantly (P < 0.05). liquid carbon dioxide. The material was then placed in an appropriate specimen tray, covered with a 30-nm layer of gold, and observed under a scanning electron microscope (Jeol JSM 25SII model) operating at 15 kv. The average number of villi/segment was obtained by counting the number of villi in 6 areas measuring 103,269 µm 2 each. Data were analyzed as a 2 (presence or absence of water) 2 (presence or absence of feed) factorial arrangement, using the general linear models procedure of SAS software [11]. RESULTS AND DISCUSSION During the first 24 h after hatching (Table 1), the treatments did not influence relative weight of the gizzard + proventriculus, yolk sac, jejunum weight, or duodenum length. supply promoted greater liver, duodenum and ileum weights and increased jejunum and ileum lengths compared with those of birds that did not receive feed. Ileum weight was also influenced by availability of water and was greater in chicks that received water (Table 1). Fortyeight hours posthatching (Table 2), the yolk sac relative weight was not affected by treatments (P > 0.05), but liver relative weight was greater, and the gizzard + proventriculus relative weight was less in birds receiving feed compared with birds without feed. A significant feed by water interaction was observed for jejunum and ileum weights and lengths 48 h posthatching (Table 3). Birds receiving feed and water had greater jejunum and ileum weights and a longer ileum length compared with those receiving only feed or only water (Table 3). Seventy-two hours posthatching (Table 4), yolk sac relative weight was not influenced by treatments, as also observed 24 and 48 h after hatching. However, liver and duodenum weights were affected by feed supply and were greater in chicks receiving feed. Gizzard + proventriculus relative weight was less in birds receiving feed. A significant (P < 0.05) feed by water interaction was observed for jejunum and ileum weights and also for duodenum, jejunum, and ileum lengths 72 h posthatching (Table 5). The birds that were supplied with feed, water, or both showed significant (P < 0.05) increases in jejunum and ileum relative weights and lengths when compared with birds that did not receive water and feed. The development of the gastrointestinal tract is directly linked to feed intake. Mitchell

7 MAIORKA ET AL.: GASTROINTESTINAL TRACT DEVELOPMENT 489 TABLE 6. Effects of deprivation of water, feed, or both after hatching on number of villi per area (103,269 µm 2 ) in different segments of the intestine 24, 48, and 72 h posthatching A Posthatching period 24 h 48 h 72 h Treatment Duodenum Jejunum Ileum Duodenum Jejunum Ileum Duodenum Jejunum Ileum With 105 ± ± ± ± 13 b 219 ± ± ± 28 b 130 ± 20 b 275 ± 58 Without 140 ± ± ± ± 30 a 256 ± ± ± 20 a 255 ± 26 a 402 ± 47 Water With 106 ± ± ± ± 36 b 227 ± ± ± 46 b 175 ± 67 b 296 ± 80 Without 138 ± ± ± ± 46 a 248 ± ± ± 39 a 210 ± 69 a 382 ± 59 Source of variation Water water NS NS NS P a,b Means within a column for each variable with no common superscript differ significantly by Tukey s test (P < 0.05). A Each value represents the mean ± SEM (n = 5).

8 490 JAPR: Field Report FIGURE 1. Scanning electron-photomicrograph of the jejunal mucosa of 1-d-old chicks. Treatments of water ad libitum and no feed (panel B) and no water and feed ad libitum treatments (panel C) showed an increase in the number of villi per area when compared with water and feed ad libitum (panel A). The water and no feed treatment (panel D) showed the highest number of villi per area ( 100). and Carlisle [12] showed that the reduction in feed intake, caused by chronic exposure of birds to high temperature (35 C), resulted in reduction in small intestine weight and length and also in changes in villus characteristics. Noy and Sklan [13] asserted that, at hatching, most of the energy and part of the protein are directed at the development of the intestine. This process may explain the reduction in gastrointestinal organ weight and length in birds subjected to deprivation of feed, water, or both observed in the present experiment. In particular, liver weight was also affected by 24 h of fasting, which demonstrated that the metabolism and development of this organ after hatching is probably associated with substrates derived from intestinal absorption. In addition, gizzard + proventriculus relative weight was significantly affected by the absence of feed for 48 and 72 h posthatching compared with those of birds fed ad libitum. Katanbaf et al. [14] observed that feed-restricted birds showed an increased gastrointestinal tract relative weight. Palo et al. [15] restricted feed of chickens from 7 to 14 d posthatching and observed that birds on restriction had higher proportional gizzard weights on d 21 but not on d 42, whereas the proportional weight of the proventriculus was significantly higher only on d 42 compared with birds fed

9 MAIORKA ET AL.: GASTROINTESTINAL TRACT DEVELOPMENT 491 TABLE 7. Interactions between feed and water treatments on number of villi per area (103,269 µ 2 ) in duodenum, jejunum, and ileum segments at 24, 48, and 72 h posthatching A Hours Water posthatching With Without Duodenum With ± 15 ay 109 ± 7 by Without 105 ± 11 az 171 ± 5 ay Jejunum With ± 7 bz 419 ± 26 by Without 352 ± 30 az 511 ± 26 ay With ± 30 ay 446 ± 32 by Without 422 ± 18 az 500 ± 8 ay Ileum Jejunum With ± 18 ay 232 ± 26 by Without 216 ± 20 az 281 ± 13 ay With ± 7 bz 315 ± 14 by Without 331 ± 13 az 399 ± 10 ay With ± 16 bz 370 ± 14 by Without 328 ± 7 az 435 ± 27 ay Ileum Ileum a,b Means within a column for each variable with no common superscript differ significantly (P < 0.05). y,z Means within a row for each variable with no common superscript differ significantly (P < 0.05). ad libitum. The results suggest that early in life broiler chickens give priority to the development of organs related to digestion (i.e., gizzard, intestine) for better feed use and future growth. The development of the intestinal mucosa primarily results from 2 associated cytological events: cell renewal (mitosis) and cell loss (extrusion). The balance between these 2 processes determines a constant turnover and the maintenance of villus size. Uni et al. [3] stated that some microvillus clumping and abnormal crypt structure occur in birds with delayed access to feed. Yamauchi et al. [10] demonstrated that, after long periods of fasting, epithelial cells begin to show large lysosomal autophagic vacuoles, which suggests that fasting can cause cell death. Cell death probably leads to increased extrusion rate and a consequent reduction in villus size, which have an important impact on intestinal area for digestion and absorption [16]. The mucosal development following delayed access to feed and water in different intestinal segments is shown in Table 6 and Figure 1. There was an increase in the number of villi per area when neither feed nor water was offered to the birds. The number of villi per area also showed a significant (P < 0.01) feed by water interaction at 24 h (duodenum, jejunum, and ileum), 48 h (jejunum and ileum), and 72 h (ileum) posthatching on the number of villi per area of chick intestinal segments (Table 7). Data analysis revealed that lack of water had the same effect as lack of feed, both showing a higher number of villi per area when compared with feed and water ad libitum treatments. This increase in the number of villi per area was due to a reduction in size; they were measured in the same area for each treatment. Tarachai and Yamauchi [17] reported that the intestinal mucosa seems to respond not only to physical stimuli, but in the presence of nutrients there is greater development of the intestinal mucosa with increased villus height, numbers of mitotic cells, and extrusion, suggesting that the primary stimuli for mucosal develop-

10 492 ment are the chemical characteristics of the nutrients. The results of this experiment also confirm that absence of physical stimuli caused by feed in the intestinal lumen and the specific need for certain nutrients, such as water, may be responsible for changes in the morphology of the intestinal mucosa. These observations have JAPR: Field Report a practical application, because usual hatchery practices result in 24- to 48-h delays between hatching and placement of birds into a poultry house. So, as demonstrated herein, delayed access to feed or water results in changes in morphological development of the intestinal tract, which could impair poultry performance [18, 19]. CONCLUSIONS AND APPLICATIONS 1. These results indicate that absence of feed and water soon after hatching negatively affect the morphological development of the intestinal mucosa, and therefore both feed and water should be offered as soon as possible after hatching to avoid delay in the development of the chick gastrointestinal tract. REFERENCES AND NOTES 1. Overton, J. and J. Shoup Fine structure of cell surface specialization in the maturing duodenal mucosa of the chick. J. Cell Biol. 21: Nitsan, Z., G. Ben-Auraham, Z. Zoref, and I. Nir Growth and development of the digestive organs and some enzymes in broiler chicks after hatching. Br. Poult. Sci. 32: Uni, Z., S. Ganot, and D. Sklan Posthatch development of mucosal function in broiler small intestine. Poult. Sci. 77: Imondi, A. R., and F. H. Bird The turnover of intestinal epithelium in the chick. Poult. Sci. 45: Baranyiova, E Influence of deutectomy, food intake and fasting on the digestive tract dimensions in chickens after hatching. Acta Vet. Brno. 41: Baranyiova, E., and J. Holman Morphological changes in the intestinal wall in fed and fasted chickens in the first week after hatching. Acta Vet. Brno. 45: Nitsan, Z., E. A. Dunnington, and P. B. Siegel Organ growth and digestive enzyme levels to fifteen days of age in lines of chickens differing in body weight. Poult. Sci. 70: Shehata, A. T., J. Lerner, and D. S. Miller Development on nutrient transporter system in chick jejunum. Am. J. Physiol. 246:G101 G Moran, E. T Digestion and absorption of carbohydrates in fowl and events through prenatal development. J. Nutr. 115: Yamauchi, K., H. Kamisoyama, and Y. Isshiki Effects of fasting and refeeding on structures of the intestinal villi and epithelial cells in White Leghorn hens. Br. Poult. Sci. 37: SAS Institute User s Guide: Statistics. SAS Institute Inc., Cary, NC. 12. Mitchell, M. A., and A. J. Carlisle The effect of chronic exposure to elevated environmental temperature on intestinal morphology and nutrient absorption in the domestic fowl (Gallus domesticus). Comp. Biochem. and Physiol. 101A: Noy, Y., and D. Sklan Energy utilization in newly hatched chicks. Poult. Sci. 78: Katanbaf, M. N., E. A. Dunnington, and P. B. Siegel Restricted feeding in early and late-feathering chickens. 3. Organ size and carcass composition. Poult. Sci. 68: Palo, P. E., J. L. Sell, F. J. Piquer, M. F. Soto-Salanove, and L. Vilaseca Effect of early nutrition restriction on broiler chickens. 1. Performance and development of the gastrointestinal tract. Poult. Sci. 74: Smith, M. W., M. A. Mitchell, and M. A. Peacock Effects of genetic selection on growth rate and intestinal structure in the domestic fowl (Gallus domesticus). Comp. Biochem. Physiol. 97A: Tarachai, P., and K. Yamauchi Effects of luminal nutrient absorption, intraluminal physical stimulation, and intravenous parenteral alimentation on the recovery responses of duodenal villus morphology following feed withdrawal in chickens. Poult. Sci. 79: Nir, I., and M. Levanom Effect of posthatch holding time on performance and on residual yolk and liver composition. Poult. Sci. 72: Pinchasov, Y., and Y. Noy Comparation of posthatch holding time and subsequent early performance of broiler chicks and turkey poults. Br. Poult. Sci. 34: ACKNOWLEDGMENTS The authors thank Fundaç o de Amparo ã Pesquisa do Estado de São Paulo (FAPESP Proc. 98/ ) and FINEP (Financiadora de Estudos e Projetos) for financial support and Electronic Microscopy Laboratory of UNESP/Jaboticabal for analyses.