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Livestock Production Science 95 (2005) 255 – 263 www.elsevier.


Stress parameters and immune response of layers under different cage floor and density conditions
E.E. OnbaYVlarT, F.T. Aksoy
Ankara University, Faculty of Veterinary Medicine, Department of Animal Science, 06110 DVYkapV, Ankara, Turkey Received 11 February 2004; received in revised form 18 November 2004; accepted 11 January 2005

Abstract The aim of the experiment was to investigate the effects of cage floor and cage density on stress parameters of laying hens. A total of 162 brown laying hens (Hyline Brown), aged 34 weeks, were used in the experiment. Compact-type battery cages, with three floors, were used. Hens were allocated as one, three or five hens in each of 18 cages to obtain three different cage density groups of 1968, 656 and 393.8 cm2 floor area per hen, respectively. The same number of cages with different cage density were allocated to three different battery floors (first floor=top, second=middle, third=bottom) systematically. Values for body weight, mortality rate, egg weight, egg production, egg quality characteristics, egg yolk cholesterol content, the levels of blood plasma corticosterone, serum glucose, total cholesterol and triglycerides, the ratio of heterophils to lymphocytes (H–L ratio), antibody titers, claw length score, foot health score, plumage score and throat skin injuries were taken as indicators of stress. The values for egg weight ( Pb0.01) at the first floor were greater than the other floor levels. The group with five hens per cage had significantly lower mean estimates ( Pb0.01) than other groups with respects to body weight ( Pb0.001), egg production ( Pb0.001), egg weight ( Pb0.001) and plumage score ( Pb0.01), while significantly higher mean estimates for egg albumen index ( Pb0.01), Haugh unit ( Pb0.01), serum glucose ( Pb0.001), and H–L ratio ( Pb0.001). Serum cholesterol was higher in cages with one hen than that with five hens, whereas plasma corticosterone was lower. Antibody titers in cages with one hen was similar to that with three or five hens; however, those with three hens had higher titers than those with five hens. Values for egg breaking strength, yolk index, egg cholesterol content, and foot health score were not affected by cage density or floor. The results suggest that the allocation of three hens per cage had no measurable effect on health and welfare. D 2005 Elsevier B.V. All rights reserved.
Keywords: Cage density; Cage floor; Laying hens; Performance; Stress parameters

1. Introduction
T Corresponding author. Tel.: +90 312 317 03 15; fax: +90 312 318 17 58. E-mail address: (E.E. OnbaYVlar). 0301-6226/$ - see front matter D 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.livprodsci.2005.01.006

Battery cage systems are the most economical for the commercial layer industry, but have negative effects on hen welfare. For this reason, there has been pressure to ban battery cage production in several

1983). Cage density is an another important factor in plumage condition. One egg from each subgroup was obtained at the end of the experiment for cholesterol measurement. There have been some controversial results on the effects of different cage systems and cage density on the stress parameters of laying hens. body weight (Hughes. yolk index.. Therefore. (1988) indicated that corticosterone is not a useful measurement of longterm stress or well-being of chickens. Animals and diets A total of 162 brown laying hens (Hyline Brown). Davami et al. including productive performance.. environment. width and length were determined. 656 and 393. production. antibody response and external appearance. changes in the number of circulating leucocytes. As a result. The ratio of heterophils to lymphocytes (H–L ratio) measures a physiological change. Davami et al. one egg was collected from each subgroup (18 eggs for each group) at 4-week intervals to determine the egg quality characteristics. The experimental period was 22 weeks. whereas the blood concentration of corticosteroid is affected by many factors (Gross and Siegel. rapid and temporary changes occur in the body initially. Cunningham and Ostrander. third=bottom) systematically. To maintain cage density. Factors causing stress include physiological factors. and diseases. 1986). Under stress. Stress is defined as the interaction between stress factors and protective reactions. Individual eggs were weighed and their shape index. to obtain three different cage density groups: 1968. Many investigations have shown that. were used in the experiment. High levels of corticosterone have been shown to be an indicator of stress. 1972). 1987) and egg production and egg weight (Robinson. 1984. 1987) reported improved plumage condition for hens housed at low cage density. Adams and Craig. For this reason. Throughout the experiment. 1987). Cunningham et al. and physical conditions. a decline in yield and resistance to diseases may occur.2. blood parameters. Materials and methods 2. Quart and Adams. and physiology.T. Hens were allocated as one. as cage density increases (Mashaly et al. 2. Eggs were collected daily and egg production was calculated as a hen–day basis.1. second=middle. and albumen height. Each cage was 48Â41Â46 cm3 (widthÂdepthÂheight). Stress in poultry has been reviewed by Siegel (1980) and Hill (1983)..256 E. in each of 18 cages. Animals under stress become ill more easily. The same number of cages. with concomitant alterations in disease resistance. 1984. were allocated to three floors (first floor=top. any dead hen in the treatment was replaced on day of death from a reserve stock maintained at the same density.. when hen density is increased. Roush et al. 1987. egg quality characteristics. 1978. egg-shell breaking strength and shell thickness were measured. 1984. 1982) decrease. OnbaYVlar.8 cm2 floor area per hen. were used. suppression of humoral immunity. 1982. Stress may include increases in circulating levels of corticosterone. the aim of this study was to assess the response of laying hen to different management systems (cage floor and cage density). nutrition. Cunningham and Gvaryahu. Egg and shell quality analyses were completed within 24 h of the eggs being collected. 1985) increases. Yolk height and width. three and five hens. Some researchers (Hill and Hunt. 1982. Quart and Adams. behaviour.E. Aksoy / Livestock Production Science 95 (2005) 255–263 countries. 2. Davami et al. aged 34 weeks. and excess medicine may be necessary to maintain health. these are followed by permanent and irreversible changes. Compacttype battery cages with three floors. albumen index and Haugh unit were calculated (Card and Nesheim.. 1987). with continuous stress. 1979. However.. By using these values. At the end. such as cage density and transport (Freeman. drug residues increase in animal products and threaten public health directly. Laying hens were examined for mortality during the experiment. Eggs . Feed and water were provided ad libitum and the diets were presented in mash form. and a decrease in growth and egg production (Mench et al. stress conditions in animals need to be examined carefully. and mortality (Koelkebeck and Cain. Stock health and welfare management are key factors in animal health and food safety. F. such as climate. Traits measured Hens were weighed individually at 34 and 56 weeks of age. Hen welfare is based on health. with different cage density. 1975. respectively. having 18 cages and 54 cages in total.

USA). Blood samples were smeared on to a glass slide for the determination of the H–L ratio.1 ml of 0. CFÂCD: CF by CD interaction. and plumage score. one hen was randomly selected from each cage and bled from the brachial vein. Aksoy / Livestock Production Science 95 (2005) 255–263 257 were weighed and boiled for 5 min. foot health. H–L ratio) and hen– day egg production to examine the time-effect with cage density and cage floor. claw length. F. A repeated-measures ANOVA was conducted on blood parameters (serum glucose. and triglyceride levels were determined using a Hitachi autoanalyzer (Hitachi. hens were individually taken out of their cage and examined for feather damage using a scoring system. Chicago. The yolk and albumen were separated and were weighed. Throat skin injuries were scored from 0 to 3 points. egg cholesterol. 2. The H–L ratios were determined by dividing the number of heterophils by the number of lymphocytes. ¨ The total leukocyte count includes heterophils. glucose. All titers were expressed as the log2 of the reciprocal of the serum dilution (Arda.. Circulating anti-SRBC antibody titers were determined by the microhemagglutination technique from samples taken at 5 days after the immunization. Plasma corticosterone levels were measured using the kits (Gamma-B 125I Corticosterone. After drying. cholesterol. Yolk cholesterol was extracted according to the AOAC (1990) method. and eosinophils. and 4 months. glucose and triglyceride levels. When a significant difference was Table 1 Effect of cage floor and cage density on body weight at the end of the experiment Body weight (g) CD1 CD3 CD5 CF1 CF2 CF3 Two-way ANOVA df CF CD CFÂCD 2 2 4 Mean squares 13. CD: cage density. and plasma corticosteroid. once on each slide.058T 12.05). Serial Number 1238-23) and its accompanying commercial kits. basophils. one hen was randomly selected from each subgroup and injected with 0. the smears were stained with May– Grunwald–Giemsa stain (Gross and Siegel. The bleeding procedure was limited to 1 min or less to minimize the influence of handling stress. Blood samples were taken in two tubes. back. . where 0=no ateromata defects on skin and 3=large defects.489 2124a 2096a 1861b 2019 2049 2013 ab: means within columns with different letters are significantly different ( Pb0. breast. 1983). All blood samples were collected at the same time in the morning and centrifuged. Code AA-13 F1) for IDS double antibody RIA technique. Claw length was measured on a 1–4 scale. where 1=extremely long and 4=normal to short claw. A score (graduated from 1=very poor plumage to 4=intact plumage) was assigned for plumage condition for each area of the body (neck. and the other had no anticoagulant for estimating cholesterol. and tail). with a Berthold LB211 gamma counter. claw length and foot health were measured. At 56 weeks of age. At 54 weeks of age. A two-way ANOVA was used to determine differences between cage density and cage floors and their interactions with respect to body weight. where 1=intact matrix and 4=severely injured matrix (Davami et al. monocytes.171. including plumage condition. wings. A three-way ANOVA. 1997). At 0. IL. Serum cholesterol. with cage density and cage floor and time as main factors. T Pb0.9% saline.670 1.E. throat skin injuries. 2. triglyceride. Plasma was frozen (À20 8C) until analyzed for corticosterone determination. External appearance traits. antibody titers. Data were tested for distribution normality and homogeneity of variance.25% suspension of sheep erythrocytes (SRBC) in 0. OnbaYVlar. CF: cage floor. was used to detect any change in egg weight or quality. using a light microscope at Â1000 magnification. Foot health and injuries to the claw fold were scored on a 1–4 scale. 1987).001.3. one contained EDTA for estimating plasma corticosterone levels and the H–L ratio. Tokyo. Antibody titers for each hen were converted to appropriate natural logarithms.T.E. lymphocytes. One hundred leucocytes were counted. Statistical analyses Statistical analyses were performed using the SPSS software package for Windows (SPSS.

7y 65.830 Between subjects (mean squares) 15.527 35. respectively. however. 1990).05.125 ab..387 20. TT Pb0. 1972) and cage density (Cunningham and Ostrander. CD: cage density.128TT 148. Roush et al. 1978. T: period. xyz: means within columns with different letters are significantly different ( Pb0. resulted in a lower body weight. and three (3.05).460 found among cage densities or among cage floors. CF: cage floor. CFÂT: CF by T interaction. F. 1982. Davami et al.0d 62. 1984. at the beginning of the experiment. Davami et al. and period on hen–day egg production Egg production (%) CD1 94.. 1970. T Pb0.05). One (5.2 85. and third floor. and Grover et al. 1998). CFÂCDÂT: CF by CD by T interaction. and five hens per cage.3.236 20.1. feather pecking was not observed. CFÂCD: CF by CD interaction. that cage floor (Adams and Jackson. 1984. versus cage density and cage floor were not statistically significant (data not shown).258 E. When a significant interaction between cage density.3z Three-way ANOVA CF CD T CFÂCD CFÂT CDÂT CFÂCDÂT df 2 2 2 4 4 4 8 Mean squares 349.. Results and discussion 3.084 1317TT 19.3 88. 1981. Hill and Hunt. TT Pb0.1a Repeated-measures ANOVA df Within-subjects contrast (mean squares) 183.5 87. 1982. 1987. from one and three to five hens/cage.56%). 3. Bonferroni’s test was used. CD: cage density. Cage floor position did not affect hen–day egg production. This is in agreement with other studies (Hughes. Roush et al.. as shown in Table 2.3b 78. zero (0. i. CF: cage floor. (1969) found that cage density had no effect on body weight of layers.712TTT 25.4b 64.70%) hens died on the first. respectively. 1998) had no effect on mortality. Feather pecking increased mortality rates in laying hens (Rodenburg et al.56%).693 40. CFÂCD: CF by CD interaction. 1987. IYcan et al.05).. 1987. which also reported low body weight with high cage density. who found that mortality increased with cage density. Aksoy / Livestock Production Science 95 (2005) 255–263 Table 3 Effect of cage floor and cage density on egg weight Egg weight (g) CD1 CD3 CD5 CF1 CF2 CF3 T1 T2 T3 63..4a 63. in the present study.00%). one (1. In contrast. Increasing cage density. second. a one-way ANOVA was used to detect differences among different conditions (SaundersDawson and Trapp.T.001..e. (1972) reported that body weight of hens maintained Table 2 Effect of cage floor. however. Adams and Craig.051TT 5334. CFÂCDÂT: CF by CD by T interaction.001.01. T Pb0. OnbaYVlar. Cunning˙ ham.. Mortality was not affected by the cage floor or density ( PN0. CFÂT: CF by T interaction. 3. xy: means within columns with different letters are significantly different ( Pb0. three (5.E. cage floor position had no effect on body weight (Table 1). 1985).4x 63. on the top cage floors was higher than those on lower floors. Cook and Dembnicki (1966) and Wayman et al. three. These results contrast with those of some researchers (Koelkebeck and Cain.8y T CFÂT CDÂT CDÂCFÂT 2 4 4 8 CF CD CFÂCD 2 2 4 abc. IYcan et al.6a 62. Grover et al. Cunningham and ˙ Gvaryahu. cage floor or period was detected.85%). T: period. 2003). CDÂT: CD by T interaction.2.4x 88. .7y 87. de. 1975..5e 60.315T 8. Increasing the cage density may stimulate feather pecking. 3. cage density. Other researchers reported similar results. Egg produc- CD3 CD5 CF1 CF2 CF3 T1 T2 T3 89. TTT Pb0. Egg production and egg weight Increasing density had a negative effect on egg production.9e 62. Body weight Differences in body weight.01.33%) hens died in the groups having one.5c 86. Mortality During the experiment.906T 773. CDÂT: CD by T interaction. and two (3. 1984.

cage floor. In contrast.06 8. Eggs in the first floor were heavier than the second or third floor.. TT Pb0.83 7.331 2. ˙ 1992.833 3. egg weight. and housing conditions. Cage density had no effect on egg shape index.512 35. having one or three hens per cage.435 2..5b 77.997TT 9.4.E. and period on some egg quality characteristics Shell thickness (Am) CD1 CD3 CD5 CF1 CF2 CF3 T1 T2 T3 T4 df CF CD T CFÂCD CFÂT CDÂT CFÂCDÂT 2 2 3 4 6 6 12 389 403 397 393d 398de 399e 381x 391x 416y 397xy Albumen index (%) 7. IYcan et al. 2004). The differences between the present study and literature reports may be due to the genotype and age of birds. maintenance. CFÂCD: CF by CD interaction. ¨ ¨ Anderson et al.1 45.48a 8. feeder space.9xz Haugh unit (%) 74.1 44. In the present experiment.355 0.8a 79. and health. whereas egg shell thickness on the third floor was larger than that on the first floor (Table 4).90b 8.6 76. Suto et al. 1998) reported that cage density had no effect on egg production or egg weight.01. while the remaining portion is divided into three equal parts for reproduction. albumen index. 1982. Cage floor. 1987. egg-shell breaking strength (data not shown). Similarly.920 ab.546 4.20 8. and body weight in groups with five hens in cages.058TT 232.. The first floor was fresher than the others.5 45. Increasing cage density from one and three to five hens per cage resulted in a lower egg weight.4bz 43. Egg weight increased significantly with time.7xy 80. Table 4 Effect of cage density. CDÂT: CD by T interaction. 2000). yolk index.E. Reports by Cook and Dembnicki (1966) and Dorminey and Arscott (1971) indicated that higher cage densities causes an increase in egg weight. xyz: means within columns with different letters are significantly different ( Pb0.T.921 40. In stress.. most of the consumed food is used to cope with unpleasant conditions (Siegel and Gross.6a 76. then increased. season. yolk index.6 77. There was an interaction between floor and density. de.12 8. egg-shell breaking strength. 1997. 1979. density.9 76. (1987) concluded that hens in lower density cages were allowed more movement within the cage.3y 74.121TT 6. None of the interactions of cage densityÂcage floor. F.00 Yolk index (%) 45.395 23. OnbaYVlar. some researchers found no differences in egg Haugh unit at different cage floors (Adams and Jackson. CFÂT: CF by T interaction. Brake and Peebles. .960 1. in egg-shell breaking strength (Wells. CD: cage density. 2000). other investigators (Koelkebeck et al.05.806 62.764 10.0 45. Food is partitioned between body functions.988 20.169 4. and time had a significant effect on egg weight (Table 3).802 12. Davami et al.413 54. and this may contributed positively to egg weight. this condition may explain the reduction in the egg production. T Pb0.. Egg yolk cholesterol content was not affected by cage density or floor position (data not shown). The heaviest eggs were detected on the first floor. Aksoy / Livestock Production Science 95 (2005) 255–263 259 tion was lower at the beginning of the study. 1970).471 5. cage densityÂperiod and cage floorÂperiod were significant for these egg quality characteristics. and Haugh unit of laying hens with time.608 290.687 2. Egg quality characteristics and egg yolk cholesterol content Egg shape index (data not shown). 10% of food ingredients consumed are used to maintain health. including maintenance.6 43. and Haugh unit of laying hens were similar for cage floor positions.344TTT 7.639 41.11a 8. Cunningham and Ostrander. 1995.001. T: period.37 8. CF: cage floor. shell thickness or yolk index.5x 76.0x Three-way ANOVA (mean squares) 24.2 44. TTT Pb0. 3.861T 5. reproduction. CFÂCDÂT: CF by CD by T interaction. The results of egg production and egg weight were similar to those reported in some studies (Robinson. Carey et al. Quart and Adams. In healthy animals. which may have resulted in a less stressful environment.652TT 89.9x 46.881 6.78 8. The values of albumen index and Haugh unit were higher in cages containing five hens than in the cages with one or three hen.322 5.05). but some changes were detected in shell thickness. and health (Siegel and Gross. 1982. growth.6y 45.

Analysis between subjects showed significant effects of cage density on plasma corticosterone. 1972).. The corticosterone levels found in the present study are in agreement with the findings of some researchers (Mashaly et al.0396 386.72ab 231b 1.00211 0. Increasing cage density. Differences in corticosterone secretion between experiments may be caused by a number of external factors.506 113. 1997). although there were no significant cage floor effect on blood parameters.62a 0.178TT 94. such as light.05). 3. cholesterol.01) and serum glucose ( Pb0. CDÂT: CD by T interaction.88x 211x 1. serum glucose. which..706 1654TT 183.. triglyceride concentrations. and H–L ratios. density.007479 1. 1988) reported that plasma corticosterone was not significant in chronic stress.1720TTT 24. and methodological factors of taking blood. (1987) and Suto et al.5230T 26.563 81.41 602. Beuving et al. cage density and period on plasma corticosterone.72 0.E.00268 0.001) concentrations and a decrease of the serum cholesterol ( Pb0. CF: cage floor. CFÂT: CF by T interaction. F. 1989). mean squares) 1996.95b 0. neural impulses come to the hypothalamus and are converted to neuro-humoral factors.74 232 1.03124 0.802 30. resulted in a significant increase of the plasma corticosterone ( Pb0.65a 221a 1. . 1983. or inherent variation (Littin and Cockrem. 1987.783T 0. triglyceride concentrations and H–L ratio Plasma corticosterone (ng/ml) CD1 CD3 CD5 CF1 CF2 CF3 T1 T2 T3 T CFÂT CDÂT CDÂCFÂT CF CD CFÂCD df 2 4 4 8 df 2 2 4 Serum glucose (mg/dl) Serum cholesterol (mg/dl) Serum triglyceride (mg/dl) Heterophils/lymphocytes 0.903 abc. (1997) reported ¨ ¨ that Haugh unit values were not affected by cage density. others (Cunningham et al. 1986a.. stimulated the adrenal for corticosterone secretion (Hill. cholesterol concentrations.77 234 1. Aksoy / Livestock Production Science 95 (2005) 255–263 Table 5 Effect of cage floor. Corticotropin-releasing factor stimulates the anterior hypophysis to secrete ACTH. However. OnbaYVlar. xyz: means within columns with different letters are significantly different ( Pb0.93b 250c 1. serum glucose. higher corticosterone levels in the 5 hens/cage groups could reflect higher stress conditions. from one to five hens per cage.0118 37.70 0. T: period.73y 236z Repeated-measures ANOVA (within-subjects 0.0168 71. Siegel. On the contrary.69y 255y 1.001.5TT 0.0130 67. Suto ¨ ¨ et al..001 171.707 573.T. temperature. Therefore. During stress conditions. CFÂCD: CF by CD interaction.4 0.68 0. TTT Pb0. such as genetic stock. 1985).382TTT 3952T 0.326 41. age.01. 1987. Craig et al.70 0.997 133a 1280 122ab 1218 114b 1089 121 1245 125 1148 122 1194 120 1086x 118 1205xy 130 1297y contrast. at varying densities. Davami et al. CFÂCDÂT: CF by CD by T interaction. serum glucose.05) concentrations. 1986.317 655.00393 0.167 0.57a 0. background. 1984.124T 0.00484 0. 2001). group size.2 Between subject (mean squares) 0.05.5. or internal factors.76 0. T Pb0.80 235 1. CD: cage density. Plasma corticosterone. or in shell thickness (Davami et al.72 0. Koelkebeck et al.. TT Pb0.465 75..260 E.329 42. and H–L ratios The repeated-measures ANOVA revealed a decrease in plasma corticosterone and an increase in serum glucose and triglyceride concentrations at the end of the experiment compared to admission levels (Table 5). cholesterol. in turn.

4. back. (1996). There were no significant differences in plumage score among different cage floors. 3.. CF: cage floor. Table 6 Effect of cage floor and cage density on antibody titers and plumage score Antibody titers (Log2) CD1 CD3 CD5 CF1 CF2 CF3 CF CD CFXCD df 2 2 4 Plumage scorea 2.22ab 16.658T 182. OnbaYVlar.895 1.42b 2.677T 0.E. This result agrees with the findings of some researchers (Quart and Adams. . The allocation of hens at 5/cage resulted in a reduction in productive performance.57a 14. Patterson and Siegel (1998) and Heckert et al. 1983.504 Increasing cage density from 3 to 5 hens/cage resulted in a significant decrease ( Pb0.25 15. Some of these differences may be due to the genetic background of the birds.. This may be possibly explained by the difference in light intensity at different cage floor positions. different cage densities or the use of different suspensions of sheep erythrocytes. Cholesterol concentrations of hens were lower at the higher density. Koelkebeck et al. CFÂCD: CF by CD interaction. F. 1987. Conclusion The study indicates that significant differences in egg weights were detected among different cage floor positions.85a 1. (1986). On the other hand. Pb0. Some strains have a greater ability to adapt to highdensity environments and this may explain the differences between experiments. antibody response to the antigen SRBC and plumage score.01. a Plumage score: 1 (very poor plumage)–4 (intact plumage) for each area of the body (neck.E. Davami et al. Siegel. The results obtained in the present study are in agreement with the findings of other researchers (Gross and Siegel. 3.56a 2. 1974. plumage score comparisons in the study of Cunningham et al.001. This condition could be explained by the elevated concentration of corticosterone in blood circulation.7. In contrast.0330 8. However. Cunningham and Gvaryahu. 1987). In contrast. are described as an important indicator of stress conditions. throat skin injuries were found only on the hens maintained on the first floor (5. The glucose concentrations found in the present study are in agreement with the findings of Lagadic et al. cage density and cage floor had no significant effect on the scores of claw length and foot health (data not shown). as was also found by Clemens et al. Simon.29 14.. 1983.01) in the antibody titers to SRBC in the present study.6% of hens). Pesti and Howarth (1983) found no changes in cholesterol levels of broilers at different population densities.T. (2002) observed that cage density treatments had no significant effect on hemaglutinin titers to SRBC. Aksoy / Livestock Production Science 95 (2005) 255–263 261 Increasing blood glucose levels. (1988) were not significantly different between the groups having different cage population sizes (4–6/ cage). as shown in Table 6. 1984). Craig et al. wings. External appearances The plumage score of hens were found to be lower in densely populated cages. Beuving et al. 1987. CD: cage density.01) than those of groups having 1 or 3 hens/cage. The poorer plumage score of densely populated cages can be caused by abrasion against cage wire or other hens.. ab: means within columns with different letters are significantly different ( Pb0. T Pb0. and tail). cage floor had no significant effect on antibody response in a two-way ANOVA. as shown in Table 6.96b 12. 1986b.6. This result agrees with the findings of Hester et al. 1985). breast.0699 3. due to the effect of glucocorticoids (Scnukro. Antibody response Cage density significantly affected antibody titers. which causes an increase in heterophil count and a decline in lymphocyte count (Hill.64 2. Interestingly. 1989).10 Two-way ANOVA (mean squares) 0. However.09 2.21 14. 1982. The ratio of heterophils to lymphocytes of the group having 5 hens/cage was higher ( Pb0. Higher density appears to cause increased levels of nervousness and feather-pecking activity. (1990).05).

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