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Behavior Patterns of Chickens to Ten Weeks of Age
MICROORGANISMS FROM GIBLETS processed turkeys. Poultry Sci. 4 3 : 934-938. Salzer, R. H., A. A. Kraft and J. C. Ayres, 196S. Effect of processing on bacteria associated with turkey giblets. Poultry Sci. 44:952-956. Stadelman, W. J., W. W. Marion and M. L. Eller, 1957. Antibiotic preservation of fresh poultry meat. Antibiotics Annual, 1956-1957; 839-842. Thatcher, F. S., and A. Loit, 1961. Comparative microflora of chlorotetracycline-treated and nontreated poultry with special reference to public health aspects. Appl. Microbiol. 9: 3945.
615
Walker, H. W., and J. C. Ayres, 1956. Incidence and kinds of microorganisms assocated with commercially dressed poultry. Appl. Microbiol. 4:345-349. Wilson, E., R. S. Paffenbarger, Jr., M. J. Foter and K. H. Lewis, 1961. Prevalence of Salmonellae in meat and poultry products. J. Infect. Diseases 109: 166-171. Woodburn, M., 1964. Incidence of salmonellae in dressed broiler-fryer chickens. Appl. Microbiol. 12: 492-495.
J. S. DAWSON AND P. B. SIEGEL Virginia Polytechnic Institute, Blacksburg, Virginia 24061 (Received for publication September 12, 1966)
A
LTHOUGH research involving young - chickens is voluminous, only meager information is available concerning their behavior prior to the age of peck-order formation. The social organization in flocks of chickens was first described by Schjelderup-Ebbe (1922) and reviews of social and individual behavior patterns were published by Guhl (1953, 1962) and WoodGush (1955). The development of behavior involves the ontogeny of various systems in an individual. Kuo (1932) observed embryonic behavior while Guhl (1958), Kruijt (1963, 1964), and Ratner (1966) showed a sequence in the appearance of post-hatching behavior patterns. The experiment reported here was designed to study the development of behavior from hatching to 70 days of age in flocks where sexes were intermingled or maintained separately. MATERIALS AND METHODS
The chickens consisted of progeny from matings of F 2 generation White Rocks which had been selected for high body weight (HW) and low body weight (LW) at eight weeks of age (Siegel, 1962). Upon
hatching, chicks were sexed and assigned to three flocks (A, B, C) by systematic randomization. Compositions of the flocks were: (A) 15 HW and 15 LW males, (B) 15 HW and 15 LW females, (C) 7 HW and 8 LW males and 8 HW and 7 LW females. Flock sizes were small to enable collection of data for individual birds. The plumage on the backs and sides of chicks were dyed to facilitate identification of individuals. Each flock was observed for 25 minutes daily for 70 consecutive days by the same individual. Observations commenced at 6:00 a.m. and the order in which the flocks were observed was randomly determined each day. Prior to the collection of data, a period of five minutes was allowed to enable birds to become adjusted to the observer. The pens were encompassed by a canvas baffle 1.2 linear m. high which restricted the birds' view of the outside of the pen to the upper portion of the observer's body which was clad in a white T-shirt. Floor space allowances were 1208 sq. cm. per chick started. Feeder space allowances on a per-chick-started basis were 6.0 linear cm. to five weeks of age and 12.2 thereaft-
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Behavior Patterns of Chickens to Ten Weeks of Age
616
J. S. DAWSON AND P. B. SIEGEL
Peck orders within each flock were determined from a combination of several methods. These consisted of dominant-subordinate relationships from direct flock observation and paired encounters between flockmates. Paired encounters were obtained by placing both individuals in an exhibition cage, 60 X 60 X 67 cm., and observing the dominant-subordinate relationship. Social behavior data were analyzed according to the method of asymptotic simultaneous confidence intervals outlined by Marks et al. (1960). The behavior of each individual was classified into three categories for analysis: initiates, recipients, and the combination of the two. Timetrend analyses and analyses of variance were made for the other behavioral traits. RESULTS AND DISCUSSION
The number of chickens resting, stretching, scratching, preening, running, frolicking, sparring and total encounters were calculated over time for all subgroups. Computer-fitted curves for the LW males
in the all-male flock are presented in Figure 1 as representative of the results obtained. Resting. Resting was most prevalent during the first few days of life after which it declined rapidly until the chicks were 3^ weeks of age. It then increased until 8 weeks of age and again declined. Resting, with the chicks being located in clusters, may be considered allelomimetic, a general type of behavior which Scott (1958) defined as a situation in which 2 or more individuals do the same thing with a mutual stimulation. This, however, would not necessarily be the sole reason for resting at very early ages. Another possible explanation is that the thermo-regulatory mechanism in chickens is not fully developed at these ages (Lamoreux and Hutt, 1939) and that grouping during resting and sleeping was primarily for warmth. No significant differences in resting were found between lines; however, differences were significant between methods of flocking and between sexes (Table 1). Means presented in Table 2 show that females rested more than males and that chicks in unisexual flocks rested more than those in the heterosexual flock. The method of flocking-sex (F-S) interaction was significant. The females rested more in the all-female flock than in the intermingled flock. The effect of flocking was inconsistent for males. If males rested less than the females and if part of the effect was allelomimetic then this interaction would be expected. Scratching. Scratching (Figure 1) increased during the first few weeks and then declined until it had almost disappeared by 7 weeks; it then reappeared at 9 weeks. Kruijt (1964) with Red Junglefowl observed no scratching on the first day and normal scratching on the third day. Our results with domestic chickens were comparable. There was a highly significant
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er. Drinking facilities consisted of a twogallon waterer. Birds were exposed to 328 luxes of continuous illumination. Characteristics measured were resting, scratching, stretching, preening, running, frolicking, sparring, threat-avoidances, peck-avoidances and fighting. A chick was considered resting when it gave the appearance of being asleep. Scraping of the litter with the claws was classified as scratching, and the extension of the limbs and/or body was termed stretching. Preening consisted of grooming of the feathers with the beak. Running was classified as swift movement about the pen, while frolicking consisted of a swift movement with the wings raised or napping. Running and frolicking were frequently mimicked by flockmates. Sparring involved two chicks jumping up and down as in fighting but with no physical contact between pairs.
617
BEHAVIOR PATTERNS ' Resting Preening Frolicking Stretching Total Encounters - Scratching Sparring Running
^Resting, scratching, and preening were average per period rather than average per bird.
-^
5
6
TIME (wks) FIG. 1. Summary of behavioral responses in time for LW $ $ in Flock A.
difference between lines for scratching flocking were not significant. The line(Table 1) with the frequency being greater method of flocking (L-F) and line-sex (Lin the HW than in the LW line (Table 3). S) interactions were not significant, but Differences between sexes and method of there was a significant F-S interaction. TABLE 1.—Analyses of variance for resting, scratching, stretching, preening, running and frolicking M.S Source of variation Between lines (L) Method of flocking (F) Between sexes (S) L-F L-S F-S L-F-S Error **P<.01. *P<.05.
Df
82
Resting
Scratching
45.4 90.1** 351.6** 14.9 51.3* 50.8* 4.2 11.6
97.2* 2.7 59.4 2.4 0.1 115.5* 14.0 21.9
Stretching 461.0** 29.6 335.2** 4.0 85.1 154.9* 73.9 34.9
Preening
Running
Frolicking
105.5 6.0 2.8 111.6 50.5 5.8 15.7 55.6
18.4 145.5* 59.7 8.5 271.7** 7.9 21.1 32.6
0.0 66.4 680.2 210.9 1,071.6* 5.7 201.6 248.9
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4
618
J. S. DAWSON AND P. B. SIEGEL TABLE 2.—Mean number of observation periods a bird was observed to be resting by line, sex and method of Males
Females Wt'd. x
Separate Intermingled Wt'd. x
HW
LW
Pooled
HW
LW
Pooled
9.9 10.6 10.1
10.8 9.0 10.2
10.4 9.7 10.2
16.7 13.4 15.6
14.0 9.9 12.7
15.4 11.8 14.2
12.9 10.8
TABLE 3.—Mean frequency of scratching1 and stretching^ by line, sex and method of flocking Method of flocking sexes Character
Scratching Stretching
1 2
Line
HW LW Wt'd. x HW LW Wt'd. x
Separate
Wt'd. x Intermingled
cfc?1
9 9
c?V
9 9'
10.0 7.6 8.8 24.1 18.6 21.4
12.7 11.4 12.1 21.4 17.3 19.4
11.5 10.2 10.8 29.6 21.1 25.1
11.1 7.4 9.4 17.5 18.1 17.7
Mean number of periods a bird was observed scratching. Mean number of times a bird was observed stretching.
9 9
Pooled
10.5 8.5 9.2 25.9 19.5 22.6
12.1 10.1 11.2 20.4 17.6 19.0
11.3 9.3 10.3 22.9 18.5 20.8
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Chicks in the all-male flock scratched less specific stimulus was observed which would frequently than those in the intermingled cause this behavior. flock, whereas chicks in the all-female flock Preening. No significant differences exscratched more frequently than those in the isted between lines, sexes or method of heterosexual flock. ' flocking for preening, as shown in Table 1. Stretching. Stretching was observed on Interactions between main variables also the first day (Figure 1). It increased rapid- were not significant. ly during the first few weeks, plateaued Preening, as shown in Figure 1, inbetween 4 and 7 weeks and then declined creased in a linear manner with age. Both to about half of its peak level at 10 weeks. lines were early feathering and juvenile As shown in Table 1, there was a highly feathers began to appear at very early ages. significant difference between sexes and It has been well documented (Jull, 1952) lines, with males stretching more than fe- that during the first 10 weeks of life juvemales and HW chicks stretching more than nile and post-juvenile plumage rapidly reLW ones (Table 3). Neither method of placed down and juvenile feathers. Since flocking nor the L-F and L-S interactions the amount of feather coverage increased were significant. The F-S interaction was during this period it would be logical to assignificant, with males in the heterosexual sume a concomitant increase in preening. flock stretching more than those in the all- Further, as Kruijt (1964) pointed out male flock, while the opposite relationship there are several forms of preening which existed for females. Although no ready ex- appear as the bird gets older. If the time planation is offered for this interaction, spent on each type of preening is additive stretching was probably not a spontaneous the total amount of preening would be exactivity since it frequently followed a peri- pected to increase with time. od when the birds were inactive. No Running. Running, an escape reaction,
619
BEHAVIOR PATTERNS
TABLE 4.—Mean number of runs by line, sex and method of flocking Method of flocking sexes Sex
cfcf 9 9
Line
HW LW HW LW Wt'd. x
- Wt'd. x Separate
Intermingled
7.6 13.1 10.6 7.8 9.8
7.2 9.4 6.9 4.8 7.1
7.5 11.8 9.3 6.8
TABLE 5.—Mean number of frolics by lines and sex Lines Sex
Wt'd. x HW
LW
22.4 23.8 23.1
29.4 16.9 23.2
25.8 20.4 23.1
however, would not necessarily frolic in the same direction as the chick that frolicked initially. Possible reasons for frolicking have been offered by Guhl (1958). He felt that this activity was caused by disturbances such as filling of feed troughs and the turning on of lights. Our results concur with this reasoning although the specific disturbances differed. For example, when the flocks reported here were frightened by unusual noises, the chicks restrained from their existing activity for a short time, perhaps exhibiting fear as described by Phillips and Siegel (1966). Then one individual would spontaneously frolic being followed by some of its flockmates. Spars. Sparring started somewhat later than frolicking, increased at a faster rate and surpassed frolicking when the chicks were 25 days of age (Figure 1). Sparring reached its peak at about 32 days and then declined at approximately the same rate. Sparring disappeared during the ninth week. These data on the emergence of frolicking and sparring are consistent with those of Guhl (1958). There were no significant differences among lines in the all-male flock with regard to initiates of spars (Table 6). LW males maintained in this flock, however, initiated significantly more spars than any of the other groups tested. The HW males maintained in the all-male flock differed significantly from the females maintained in flocks where sexes were separated. Differences between female groups, whether intermingled with males or maintained separately, were not significantly different.
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was observed in all groups during the first week. This substantiated the findings of Guhl (1958) who noted that running occurred very early in life. Running (Figure 1) decreased with time and was probably replaced by other escape responses. Although no significant differences were found between sexes and lines, method of flocking was significant (Table 1). Chicks reared in flocks where sexes were maintained separately ran more frequently than those reared in the sex-intermingled flock (Table 4). This suggests that the mixing of sexes reduced this type of behavior. The only significant interaction was the L-S interaction, which resulted from LW males running more frequently than the HW males and the HW females running more frequently than the LW ones. Frolicking. Frolicking (Figure 1) increased until about 4 weeks of age, declined rapidly and disappeared at about 9 weeks. The analysis presented in Table 1 shows no significant differences between sexes, lines and method of flocking for this characteristic. The L-S interaction was the only significant interaction. This interaction resulted from LW males frolicking more frequently than HW ones and HW females frolicking more than LW ones (Table 5). Frolicking appeared to occur spontaneously. One chick would frolic and almost immediately others would mimic the pattern. Those that mimicked the pattern,
620
J. S. DAWSON AND P. B. SIEGEL
TABLE 6.—Comparisons between lines within sexes within method offlockingfor spars and total encounters Separate Behav.
Role
Intermingled
&
1
9 9
9 9
cftf
HW
LW
HW
LW
HW
LW
HW
LW
Spars
Init.12
Red. 3 Comb.
76"° 84"" 160"
140° 131" 271°
28»" 19" 47"
8" 16" 24"
37"" 30" 66"
33"" 33" 67"
25" 24" 49"
10" 14" 24"
Total Enco.
Init. Red. Comb.
72" 50"" 122"
90" 112" 202°
18" 14" 32"
4" 8» 12"
23" 8" 31"
46»" 32" 78""
12" 17" 29"
1" 15" 16"
Differences between lines and sexes within the intermingled flock were not significant for recipients of spars, nor did HW and LW birds maintained in the allfemale flock differ significantly from the groups in the intermingled flock. Although the same was true for HW males in the allmale group, the LW cockerels in this flock, were significantly different from all of the other groups, except their HW flockmates. When the initiates and recipients of spars were combined, clear-cut differences became apparent. Males from the LW line in the all-cockerel flock sparred significantly more than birds in any of the other groups. The HW males, while sparring significantly less than their LW flockmates, sparred more than any of the birds in the other groups. No significant differences were noted between the other groups. Encounters. Sparring apparently lead to encounters between flockmates. These encounters exhibited definite aggressiveness and submissiveness between the participants and included peck-avoidances, threatavoidances and fights. Agonistic behavior came into focus between the second and third week, rose rapidly and replaced sparring during the seventh week (Figure 1). Encounters peaked at about 8 weeks of age and with the pecking relationship between
birds becoming unindirectional due to the establishment of peck-rights, the frequency of encounters declined. As shown by total encounters in Table 6, agonistic behavior was greatest in the allmale flock. Although cockerels from the two lines maintained in an all-male flock did not differ significantly from each other in the initiation of agonistic behavior, they differed significantly from all others, except from the LW males in the sex-intermingled flock. Differences between females in the all-female flock and in the sex-intermingled flocks were not significant. Regarding recipients of agonistic behavior, values for the LW cockerels in the allmale flock were significantly greater than for any other group with the exception of their HW male flockmates. This latter group, however, did not differ significantly from any of the other groups. When initiates and recipients were combined, the total number of encounters for each line of males in the all-male flocks was significantly greater than that of any other group with the exception of the males from the LW line in the intermingled flock. This latter group, however, did not differ significantly from the intermingled males in the HW line or from any of the female groups.
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Any two numbers on the horizontal which do not have the same superscript are significantly different at 1the 5 percent level. Initiates. 2 Recipients. 3 Combined.
BEHAVIOR PATTERNS
621
TABLE 7.—Peck-orders listed in descending order for each flock1 Flock A
Flock B
(All c?
(All 9 9 )
1 2
HI H7 H13, L22 H12 H8, H14 Hll L28 H5 L21 H3, H9 HIS, L17 L25, L30 L27 L23, L26 L19
(&&Y HI Lll H5, H6, H7 H4 H8 H9, L12, LIS L10 L3, L13 H2
(9 9)2 H19, H21 H20 H23 H22, L26 L27 H18 H17 H16 L29 L28 L24
Letter denotes line, high (H) or low (L); number denotes individual within line. Significant U values.
Mann-Whitney U Tests (Siegel, 1956) were conducted to determine if there were significant differences between lines for rank in the peck order. Values of U were computed on a within-sex basis in the flock where sexes were intermingled, whereas in sex-separated flocks a U value was obtained for each flock. Birds from the HW line ranked significantly higher in the social hierarchy than those from the LW line in three comparisons and approached significance in a fourth (Table 7). Although LW birds were involved in more encounters, they were not in the dominant positions when peck-orders were formed. Comparisons of these lines in paired encounters (Siegel and Siegel, 1963) have also shown that males in the HW line won significantly more paired encounters than those in the LW line. Since rank in the social order can influence production traits (Guhl, 1962) intermingling of genetically dissimilar stocks could bias results in certain experiments. SUMMARY
The development of behavioral patterns was studied in male and female chickens
from lines maintained under different flocking situations. Resting, stretching, scratching, preening, running, frolicking and sparring were observed during the first week of age and their frequency measured to 10 weeks of age. Aggressive behavior was observed to replace sparring which supplanted frolicking. Encounters among birds increased with time until peck-orders were established. Birds in the HW line had significantly higher ranks in the social hierarchy than those in the LW lines in 3 of 4 peck-orders. Agonistic behavior was significantly greater in the all-male flock than in either the all-female flock or the flock where sexes were intermingled. Chickens in the all-male flock had consistently greater numbers of total encounters than those in other flocks. Birds from the LW line maintained in allmale flocks had a greater number of total encounters than any other group tested, however, when peck-orders were established birds from this group ranked in the lower social positions. REFERENCES Guhl, A. M., 1953. Social behavior of the domestic fowl. Kansas Agr. Expt. Sta. Tech. Bull, 73.
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H12 H6, H15 H4 Hll, HI L28, L30 H9 H5 L16, L23 H2, H4, L2S H10, L26 H8, H13, L18 L22 L29 L27, H3 H7 L24
Flock C 2
622
J. S. DAWSON AND P. B. SIEGEL Phillips, R. E., and P. B. Siegel, 1966. Development of fear in chicks of two closely related genetic lines. Anim. Behav. 14: 84-88. Ratner, S. C , 1966. Comparisons between behavioral development of normal and isolated domestic fowl. Anim. Behav. 13 : 497-503. Scott, J. P., 1958. Animal Behavior. University of Chicago Press, Chicago. Schjelderup-Ebbe, T., 1922. Beitrage zur Socialpsychologie des Haushuhn. Zeitschrift. Psychol. 88: 225-252. Siegel, P. B., 1962. Selection for body weight at eight weeks of age. Short term response and heritabilities. Poultry Sci. 4 1 : 954-962. Siegel, P. B., and H. S. Siegel, 1963. The correlated response of relative aggressiveness to selection for body weight and breast angle in chickens. Poultry Sci. 42 : 1208-1211. Siegel, S., 1956. Non-Parametric Statistics. McGraw-Hill Book Co., Inc., New York. Wood-Gush, D. G. M., 1955. The behavior of the domestic chicken: A review of literature. Brit. J. Anim. Behav. 3 : 81-110.
The Value of Cassava Root Meal for Chicks 12 F. Q. ENEIQUEZ3 AND ERNEST Ross Department of Animal Sciences, University of Hawaii, Honolulu, Hawaii 96822 (Received for publication September 13, 1966)
C
ASSAVA (Manihot esculenta Crantz = M. utillssima Pokl) is one of the most productive root crops in tropical areas in terms of dry matter yield per acre. Seemanthani (1962) reported yields of over 15.9 and 28.4 tons per hectare for two improved varieties. An experimental plot of cassava at the University of Hawaii yielded at the rate of almost 50 tons per hectare. The ease of propagation from cuttings and the econ1 Published with the approval of the Director of the Hawaii Agricultural Experiment Station as Technical Paper No. 833. 2 From a thesis submitted to the faculty of the University of Hawaii by the senior author in partial fulfillment of the M.S. degree. 3 Present address: San Miguel Poultry and Livestock Feed Plant, Quezon City, Philippines.
omy of production make cassava a cheap and valuable source of carbohydrate. Literature on the use of cassava products in chick rations is meagre and appears to be limited to a few preliminary studies. Taboyoyong (1935) noted poorer chick growth when half or all of the rice bran in the ration was replaced by cassava refuse meal. Squibb and Wyld (1951), using a by-product residue of cassava roots (yuca) obtained satisfactory chick growth in 4 out of 7 experiments. Torres (1957-58) suggested the presence of a toxic factor in cassava root meal to explain the unsatisfactory feed consumption and high mortality observed when 30 percent of cassava root meal replaced wheat bran and middlings in the chick diet. The presence of linamarin, a
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Guhl, A. M., 1958. The development of social organization in the domestic chick. Anim. Behav. 6: 92-110. Guhl, A. M., 1962. The behavior of chickens, in The Behavior of Domestic Animal. Bailliere, Tindall & Cox, London. Jull, M. A., 1952. Poultry Breeding. John Wiley & Sons, Inc., New York. Kruijt, J. P., 1963. Ontogeny of social behavior patterns in jungle-fowl. Proc. 16th Inter. Cong. Zool. 4 : 367-370. Kruijt, J. P., 1964. Ontogeny of social behavior in Burmese Red Junglefowl. Suppl. X X I Tijdeschrift Behaviour. Kuo, Z. Y., 1932. Ontogeny of embryonic behavior in Aves. J. Expt. Zool. 6 1 : 395^129. Lamoreux, W. F. and F. B. Hutt, 1939. Variability of body temperature in the normal chick. Poultry Sci. 18: 70-75. Marks, H. L., P. B. Siegel and C. Y. Kramer, 1960. Effect of comb and wattle removal on the social organization of mixed flocks of chickens. Anim. Behav. 6: 192-196.
615
Walker, H. W., and J. C. Ayres, 1956. Incidence and kinds of microorganisms assocated with commercially dressed poultry. Appl. Microbiol. 4:345-349. Wilson, E., R. S. Paffenbarger, Jr., M. J. Foter and K. H. Lewis, 1961. Prevalence of Salmonellae in meat and poultry products. J. Infect. Diseases 109: 166-171. Woodburn, M., 1964. Incidence of salmonellae in dressed broiler-fryer chickens. Appl. Microbiol. 12: 492-495.
J. S. DAWSON AND P. B. SIEGEL Virginia Polytechnic Institute, Blacksburg, Virginia 24061 (Received for publication September 12, 1966)
A
LTHOUGH research involving young - chickens is voluminous, only meager information is available concerning their behavior prior to the age of peck-order formation. The social organization in flocks of chickens was first described by Schjelderup-Ebbe (1922) and reviews of social and individual behavior patterns were published by Guhl (1953, 1962) and WoodGush (1955). The development of behavior involves the ontogeny of various systems in an individual. Kuo (1932) observed embryonic behavior while Guhl (1958), Kruijt (1963, 1964), and Ratner (1966) showed a sequence in the appearance of post-hatching behavior patterns. The experiment reported here was designed to study the development of behavior from hatching to 70 days of age in flocks where sexes were intermingled or maintained separately. MATERIALS AND METHODS
The chickens consisted of progeny from matings of F 2 generation White Rocks which had been selected for high body weight (HW) and low body weight (LW) at eight weeks of age (Siegel, 1962). Upon
hatching, chicks were sexed and assigned to three flocks (A, B, C) by systematic randomization. Compositions of the flocks were: (A) 15 HW and 15 LW males, (B) 15 HW and 15 LW females, (C) 7 HW and 8 LW males and 8 HW and 7 LW females. Flock sizes were small to enable collection of data for individual birds. The plumage on the backs and sides of chicks were dyed to facilitate identification of individuals. Each flock was observed for 25 minutes daily for 70 consecutive days by the same individual. Observations commenced at 6:00 a.m. and the order in which the flocks were observed was randomly determined each day. Prior to the collection of data, a period of five minutes was allowed to enable birds to become adjusted to the observer. The pens were encompassed by a canvas baffle 1.2 linear m. high which restricted the birds' view of the outside of the pen to the upper portion of the observer's body which was clad in a white T-shirt. Floor space allowances were 1208 sq. cm. per chick started. Feeder space allowances on a per-chick-started basis were 6.0 linear cm. to five weeks of age and 12.2 thereaft-
Downloaded from http://ps.oxfordjournals.org/ by guest on May 7, 2015
Behavior Patterns of Chickens to Ten Weeks of Age
616
J. S. DAWSON AND P. B. SIEGEL
Peck orders within each flock were determined from a combination of several methods. These consisted of dominant-subordinate relationships from direct flock observation and paired encounters between flockmates. Paired encounters were obtained by placing both individuals in an exhibition cage, 60 X 60 X 67 cm., and observing the dominant-subordinate relationship. Social behavior data were analyzed according to the method of asymptotic simultaneous confidence intervals outlined by Marks et al. (1960). The behavior of each individual was classified into three categories for analysis: initiates, recipients, and the combination of the two. Timetrend analyses and analyses of variance were made for the other behavioral traits. RESULTS AND DISCUSSION
The number of chickens resting, stretching, scratching, preening, running, frolicking, sparring and total encounters were calculated over time for all subgroups. Computer-fitted curves for the LW males
in the all-male flock are presented in Figure 1 as representative of the results obtained. Resting. Resting was most prevalent during the first few days of life after which it declined rapidly until the chicks were 3^ weeks of age. It then increased until 8 weeks of age and again declined. Resting, with the chicks being located in clusters, may be considered allelomimetic, a general type of behavior which Scott (1958) defined as a situation in which 2 or more individuals do the same thing with a mutual stimulation. This, however, would not necessarily be the sole reason for resting at very early ages. Another possible explanation is that the thermo-regulatory mechanism in chickens is not fully developed at these ages (Lamoreux and Hutt, 1939) and that grouping during resting and sleeping was primarily for warmth. No significant differences in resting were found between lines; however, differences were significant between methods of flocking and between sexes (Table 1). Means presented in Table 2 show that females rested more than males and that chicks in unisexual flocks rested more than those in the heterosexual flock. The method of flocking-sex (F-S) interaction was significant. The females rested more in the all-female flock than in the intermingled flock. The effect of flocking was inconsistent for males. If males rested less than the females and if part of the effect was allelomimetic then this interaction would be expected. Scratching. Scratching (Figure 1) increased during the first few weeks and then declined until it had almost disappeared by 7 weeks; it then reappeared at 9 weeks. Kruijt (1964) with Red Junglefowl observed no scratching on the first day and normal scratching on the third day. Our results with domestic chickens were comparable. There was a highly significant
Downloaded from http://ps.oxfordjournals.org/ by guest on May 7, 2015
er. Drinking facilities consisted of a twogallon waterer. Birds were exposed to 328 luxes of continuous illumination. Characteristics measured were resting, scratching, stretching, preening, running, frolicking, sparring, threat-avoidances, peck-avoidances and fighting. A chick was considered resting when it gave the appearance of being asleep. Scraping of the litter with the claws was classified as scratching, and the extension of the limbs and/or body was termed stretching. Preening consisted of grooming of the feathers with the beak. Running was classified as swift movement about the pen, while frolicking consisted of a swift movement with the wings raised or napping. Running and frolicking were frequently mimicked by flockmates. Sparring involved two chicks jumping up and down as in fighting but with no physical contact between pairs.
617
BEHAVIOR PATTERNS ' Resting Preening Frolicking Stretching Total Encounters - Scratching Sparring Running
^Resting, scratching, and preening were average per period rather than average per bird.
-^
5
6
TIME (wks) FIG. 1. Summary of behavioral responses in time for LW $ $ in Flock A.
difference between lines for scratching flocking were not significant. The line(Table 1) with the frequency being greater method of flocking (L-F) and line-sex (Lin the HW than in the LW line (Table 3). S) interactions were not significant, but Differences between sexes and method of there was a significant F-S interaction. TABLE 1.—Analyses of variance for resting, scratching, stretching, preening, running and frolicking M.S Source of variation Between lines (L) Method of flocking (F) Between sexes (S) L-F L-S F-S L-F-S Error **P<.01. *P<.05.
Df
82
Resting
Scratching
45.4 90.1** 351.6** 14.9 51.3* 50.8* 4.2 11.6
97.2* 2.7 59.4 2.4 0.1 115.5* 14.0 21.9
Stretching 461.0** 29.6 335.2** 4.0 85.1 154.9* 73.9 34.9
Preening
Running
Frolicking
105.5 6.0 2.8 111.6 50.5 5.8 15.7 55.6
18.4 145.5* 59.7 8.5 271.7** 7.9 21.1 32.6
0.0 66.4 680.2 210.9 1,071.6* 5.7 201.6 248.9
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4
618
J. S. DAWSON AND P. B. SIEGEL TABLE 2.—Mean number of observation periods a bird was observed to be resting by line, sex and method of Males
Females Wt'd. x
Separate Intermingled Wt'd. x
HW
LW
Pooled
HW
LW
Pooled
9.9 10.6 10.1
10.8 9.0 10.2
10.4 9.7 10.2
16.7 13.4 15.6
14.0 9.9 12.7
15.4 11.8 14.2
12.9 10.8
TABLE 3.—Mean frequency of scratching1 and stretching^ by line, sex and method of flocking Method of flocking sexes Character
Scratching Stretching
1 2
Line
HW LW Wt'd. x HW LW Wt'd. x
Separate
Wt'd. x Intermingled
cfc?1
9 9
c?V
9 9'
10.0 7.6 8.8 24.1 18.6 21.4
12.7 11.4 12.1 21.4 17.3 19.4
11.5 10.2 10.8 29.6 21.1 25.1
11.1 7.4 9.4 17.5 18.1 17.7
Mean number of periods a bird was observed scratching. Mean number of times a bird was observed stretching.
9 9
Pooled
10.5 8.5 9.2 25.9 19.5 22.6
12.1 10.1 11.2 20.4 17.6 19.0
11.3 9.3 10.3 22.9 18.5 20.8
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Chicks in the all-male flock scratched less specific stimulus was observed which would frequently than those in the intermingled cause this behavior. flock, whereas chicks in the all-female flock Preening. No significant differences exscratched more frequently than those in the isted between lines, sexes or method of heterosexual flock. ' flocking for preening, as shown in Table 1. Stretching. Stretching was observed on Interactions between main variables also the first day (Figure 1). It increased rapid- were not significant. ly during the first few weeks, plateaued Preening, as shown in Figure 1, inbetween 4 and 7 weeks and then declined creased in a linear manner with age. Both to about half of its peak level at 10 weeks. lines were early feathering and juvenile As shown in Table 1, there was a highly feathers began to appear at very early ages. significant difference between sexes and It has been well documented (Jull, 1952) lines, with males stretching more than fe- that during the first 10 weeks of life juvemales and HW chicks stretching more than nile and post-juvenile plumage rapidly reLW ones (Table 3). Neither method of placed down and juvenile feathers. Since flocking nor the L-F and L-S interactions the amount of feather coverage increased were significant. The F-S interaction was during this period it would be logical to assignificant, with males in the heterosexual sume a concomitant increase in preening. flock stretching more than those in the all- Further, as Kruijt (1964) pointed out male flock, while the opposite relationship there are several forms of preening which existed for females. Although no ready ex- appear as the bird gets older. If the time planation is offered for this interaction, spent on each type of preening is additive stretching was probably not a spontaneous the total amount of preening would be exactivity since it frequently followed a peri- pected to increase with time. od when the birds were inactive. No Running. Running, an escape reaction,
619
BEHAVIOR PATTERNS
TABLE 4.—Mean number of runs by line, sex and method of flocking Method of flocking sexes Sex
cfcf 9 9
Line
HW LW HW LW Wt'd. x
- Wt'd. x Separate
Intermingled
7.6 13.1 10.6 7.8 9.8
7.2 9.4 6.9 4.8 7.1
7.5 11.8 9.3 6.8
TABLE 5.—Mean number of frolics by lines and sex Lines Sex
Wt'd. x HW
LW
22.4 23.8 23.1
29.4 16.9 23.2
25.8 20.4 23.1
however, would not necessarily frolic in the same direction as the chick that frolicked initially. Possible reasons for frolicking have been offered by Guhl (1958). He felt that this activity was caused by disturbances such as filling of feed troughs and the turning on of lights. Our results concur with this reasoning although the specific disturbances differed. For example, when the flocks reported here were frightened by unusual noises, the chicks restrained from their existing activity for a short time, perhaps exhibiting fear as described by Phillips and Siegel (1966). Then one individual would spontaneously frolic being followed by some of its flockmates. Spars. Sparring started somewhat later than frolicking, increased at a faster rate and surpassed frolicking when the chicks were 25 days of age (Figure 1). Sparring reached its peak at about 32 days and then declined at approximately the same rate. Sparring disappeared during the ninth week. These data on the emergence of frolicking and sparring are consistent with those of Guhl (1958). There were no significant differences among lines in the all-male flock with regard to initiates of spars (Table 6). LW males maintained in this flock, however, initiated significantly more spars than any of the other groups tested. The HW males maintained in the all-male flock differed significantly from the females maintained in flocks where sexes were separated. Differences between female groups, whether intermingled with males or maintained separately, were not significantly different.
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was observed in all groups during the first week. This substantiated the findings of Guhl (1958) who noted that running occurred very early in life. Running (Figure 1) decreased with time and was probably replaced by other escape responses. Although no significant differences were found between sexes and lines, method of flocking was significant (Table 1). Chicks reared in flocks where sexes were maintained separately ran more frequently than those reared in the sex-intermingled flock (Table 4). This suggests that the mixing of sexes reduced this type of behavior. The only significant interaction was the L-S interaction, which resulted from LW males running more frequently than the HW males and the HW females running more frequently than the LW ones. Frolicking. Frolicking (Figure 1) increased until about 4 weeks of age, declined rapidly and disappeared at about 9 weeks. The analysis presented in Table 1 shows no significant differences between sexes, lines and method of flocking for this characteristic. The L-S interaction was the only significant interaction. This interaction resulted from LW males frolicking more frequently than HW ones and HW females frolicking more than LW ones (Table 5). Frolicking appeared to occur spontaneously. One chick would frolic and almost immediately others would mimic the pattern. Those that mimicked the pattern,
620
J. S. DAWSON AND P. B. SIEGEL
TABLE 6.—Comparisons between lines within sexes within method offlockingfor spars and total encounters Separate Behav.
Role
Intermingled
&
1
9 9
9 9
cftf
HW
LW
HW
LW
HW
LW
HW
LW
Spars
Init.12
Red. 3 Comb.
76"° 84"" 160"
140° 131" 271°
28»" 19" 47"
8" 16" 24"
37"" 30" 66"
33"" 33" 67"
25" 24" 49"
10" 14" 24"
Total Enco.
Init. Red. Comb.
72" 50"" 122"
90" 112" 202°
18" 14" 32"
4" 8» 12"
23" 8" 31"
46»" 32" 78""
12" 17" 29"
1" 15" 16"
Differences between lines and sexes within the intermingled flock were not significant for recipients of spars, nor did HW and LW birds maintained in the allfemale flock differ significantly from the groups in the intermingled flock. Although the same was true for HW males in the allmale group, the LW cockerels in this flock, were significantly different from all of the other groups, except their HW flockmates. When the initiates and recipients of spars were combined, clear-cut differences became apparent. Males from the LW line in the all-cockerel flock sparred significantly more than birds in any of the other groups. The HW males, while sparring significantly less than their LW flockmates, sparred more than any of the birds in the other groups. No significant differences were noted between the other groups. Encounters. Sparring apparently lead to encounters between flockmates. These encounters exhibited definite aggressiveness and submissiveness between the participants and included peck-avoidances, threatavoidances and fights. Agonistic behavior came into focus between the second and third week, rose rapidly and replaced sparring during the seventh week (Figure 1). Encounters peaked at about 8 weeks of age and with the pecking relationship between
birds becoming unindirectional due to the establishment of peck-rights, the frequency of encounters declined. As shown by total encounters in Table 6, agonistic behavior was greatest in the allmale flock. Although cockerels from the two lines maintained in an all-male flock did not differ significantly from each other in the initiation of agonistic behavior, they differed significantly from all others, except from the LW males in the sex-intermingled flock. Differences between females in the all-female flock and in the sex-intermingled flocks were not significant. Regarding recipients of agonistic behavior, values for the LW cockerels in the allmale flock were significantly greater than for any other group with the exception of their HW male flockmates. This latter group, however, did not differ significantly from any of the other groups. When initiates and recipients were combined, the total number of encounters for each line of males in the all-male flocks was significantly greater than that of any other group with the exception of the males from the LW line in the intermingled flock. This latter group, however, did not differ significantly from the intermingled males in the HW line or from any of the female groups.
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Any two numbers on the horizontal which do not have the same superscript are significantly different at 1the 5 percent level. Initiates. 2 Recipients. 3 Combined.
BEHAVIOR PATTERNS
621
TABLE 7.—Peck-orders listed in descending order for each flock1 Flock A
Flock B
(All c?
(All 9 9 )
1 2
HI H7 H13, L22 H12 H8, H14 Hll L28 H5 L21 H3, H9 HIS, L17 L25, L30 L27 L23, L26 L19
(&&Y HI Lll H5, H6, H7 H4 H8 H9, L12, LIS L10 L3, L13 H2
(9 9)2 H19, H21 H20 H23 H22, L26 L27 H18 H17 H16 L29 L28 L24
Letter denotes line, high (H) or low (L); number denotes individual within line. Significant U values.
Mann-Whitney U Tests (Siegel, 1956) were conducted to determine if there were significant differences between lines for rank in the peck order. Values of U were computed on a within-sex basis in the flock where sexes were intermingled, whereas in sex-separated flocks a U value was obtained for each flock. Birds from the HW line ranked significantly higher in the social hierarchy than those from the LW line in three comparisons and approached significance in a fourth (Table 7). Although LW birds were involved in more encounters, they were not in the dominant positions when peck-orders were formed. Comparisons of these lines in paired encounters (Siegel and Siegel, 1963) have also shown that males in the HW line won significantly more paired encounters than those in the LW line. Since rank in the social order can influence production traits (Guhl, 1962) intermingling of genetically dissimilar stocks could bias results in certain experiments. SUMMARY
The development of behavioral patterns was studied in male and female chickens
from lines maintained under different flocking situations. Resting, stretching, scratching, preening, running, frolicking and sparring were observed during the first week of age and their frequency measured to 10 weeks of age. Aggressive behavior was observed to replace sparring which supplanted frolicking. Encounters among birds increased with time until peck-orders were established. Birds in the HW line had significantly higher ranks in the social hierarchy than those in the LW lines in 3 of 4 peck-orders. Agonistic behavior was significantly greater in the all-male flock than in either the all-female flock or the flock where sexes were intermingled. Chickens in the all-male flock had consistently greater numbers of total encounters than those in other flocks. Birds from the LW line maintained in allmale flocks had a greater number of total encounters than any other group tested, however, when peck-orders were established birds from this group ranked in the lower social positions. REFERENCES Guhl, A. M., 1953. Social behavior of the domestic fowl. Kansas Agr. Expt. Sta. Tech. Bull, 73.
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H12 H6, H15 H4 Hll, HI L28, L30 H9 H5 L16, L23 H2, H4, L2S H10, L26 H8, H13, L18 L22 L29 L27, H3 H7 L24
Flock C 2
622
J. S. DAWSON AND P. B. SIEGEL Phillips, R. E., and P. B. Siegel, 1966. Development of fear in chicks of two closely related genetic lines. Anim. Behav. 14: 84-88. Ratner, S. C , 1966. Comparisons between behavioral development of normal and isolated domestic fowl. Anim. Behav. 13 : 497-503. Scott, J. P., 1958. Animal Behavior. University of Chicago Press, Chicago. Schjelderup-Ebbe, T., 1922. Beitrage zur Socialpsychologie des Haushuhn. Zeitschrift. Psychol. 88: 225-252. Siegel, P. B., 1962. Selection for body weight at eight weeks of age. Short term response and heritabilities. Poultry Sci. 4 1 : 954-962. Siegel, P. B., and H. S. Siegel, 1963. The correlated response of relative aggressiveness to selection for body weight and breast angle in chickens. Poultry Sci. 42 : 1208-1211. Siegel, S., 1956. Non-Parametric Statistics. McGraw-Hill Book Co., Inc., New York. Wood-Gush, D. G. M., 1955. The behavior of the domestic chicken: A review of literature. Brit. J. Anim. Behav. 3 : 81-110.
The Value of Cassava Root Meal for Chicks 12 F. Q. ENEIQUEZ3 AND ERNEST Ross Department of Animal Sciences, University of Hawaii, Honolulu, Hawaii 96822 (Received for publication September 13, 1966)
C
ASSAVA (Manihot esculenta Crantz = M. utillssima Pokl) is one of the most productive root crops in tropical areas in terms of dry matter yield per acre. Seemanthani (1962) reported yields of over 15.9 and 28.4 tons per hectare for two improved varieties. An experimental plot of cassava at the University of Hawaii yielded at the rate of almost 50 tons per hectare. The ease of propagation from cuttings and the econ1 Published with the approval of the Director of the Hawaii Agricultural Experiment Station as Technical Paper No. 833. 2 From a thesis submitted to the faculty of the University of Hawaii by the senior author in partial fulfillment of the M.S. degree. 3 Present address: San Miguel Poultry and Livestock Feed Plant, Quezon City, Philippines.
omy of production make cassava a cheap and valuable source of carbohydrate. Literature on the use of cassava products in chick rations is meagre and appears to be limited to a few preliminary studies. Taboyoyong (1935) noted poorer chick growth when half or all of the rice bran in the ration was replaced by cassava refuse meal. Squibb and Wyld (1951), using a by-product residue of cassava roots (yuca) obtained satisfactory chick growth in 4 out of 7 experiments. Torres (1957-58) suggested the presence of a toxic factor in cassava root meal to explain the unsatisfactory feed consumption and high mortality observed when 30 percent of cassava root meal replaced wheat bran and middlings in the chick diet. The presence of linamarin, a
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Guhl, A. M., 1958. The development of social organization in the domestic chick. Anim. Behav. 6: 92-110. Guhl, A. M., 1962. The behavior of chickens, in The Behavior of Domestic Animal. Bailliere, Tindall & Cox, London. Jull, M. A., 1952. Poultry Breeding. John Wiley & Sons, Inc., New York. Kruijt, J. P., 1963. Ontogeny of social behavior patterns in jungle-fowl. Proc. 16th Inter. Cong. Zool. 4 : 367-370. Kruijt, J. P., 1964. Ontogeny of social behavior in Burmese Red Junglefowl. Suppl. X X I Tijdeschrift Behaviour. Kuo, Z. Y., 1932. Ontogeny of embryonic behavior in Aves. J. Expt. Zool. 6 1 : 395^129. Lamoreux, W. F. and F. B. Hutt, 1939. Variability of body temperature in the normal chick. Poultry Sci. 18: 70-75. Marks, H. L., P. B. Siegel and C. Y. Kramer, 1960. Effect of comb and wattle removal on the social organization of mixed flocks of chickens. Anim. Behav. 6: 192-196.