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IJEB 56(11) 781-794.Pdf Indian Journal of Experimental Biology Vol. 56, November 2018, pp. 781-794 Inter-relationship of behaviour, faecal testosterone levels and glandular volatiles in determination of dominance in male Blackbuck Thangavel Rajagopal1,2, Ponnirul Ponmanickam3, Arunachalam Chinnathambi4, Parasuraman Padmanabhan5, Balazs Gulyas5 & Govindaraju Archunan1* 1Pheromone Technology Lab, Department of Animal Science, Bharathidasan University, Tiruchirappalli-620 024, Tamil Nadu, India 2Department of Zoology and Microbiology, Thiagarajar College (Autonomous), Madurai-625 009, Tamil Nadu, India 3Department of Zoology, Ayya Nadar Janaki Ammal College (Autonomous), Sivakasi 626 124, Tamil Nadu, India 4Department of Botany and Microbiology, College of Science, King Saud University, Riyadh, Saudi Arabia-11451 5Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore-637553 Received 28 March 2017; revised 25 November 2017 The social hierarchy of blackbuck plays a crucial role in mate selection and establishment of hierarchy in order to maintain successful reproduction. The factors that influence dominancy have not been yet investigated in the Indian male blackbuck. Here, we investigated the interrelationships between behaviours (aggressive and scent marking), chemical profiles of preorbital gland secretion and faecal testosterone levels in male blackbuck with special reference to dominance hierarchy. The frequency of aggressive behaviour, preorbital gland scent marking behaviour and faecal testosterone level were significantly higher (P <0.001) in the dominant males than the other males. Among the 43 major volatile compounds identified in the pre-orbital gland posting of dominant and subordinate male Blackbucks, four compounds viz., 2-methyl propanoic acid (I), 2-methyl-4-heptanone (II), 2,7-dimethyl-1-octanol (III) and 1,15-pentadecanediol (IV) were present only in the preorbital gland post of the dominant male during the hierarchy period. The results suggest that male blackbuck preorbital gland marking behaviour is meant to defend the home range or territorial region, and the scent odour suppresses aggression, scent marking, scent production and territorial patrolling of subordinate males. The behavioural expression, level of testosterone and volatile chemical profiles are closely interlinked towards establishment of dominance in the blackbuck. This investigation is the first of its kind to detect establishment of dominance hierarchy based on identification of volatile chemical compounds of preorbital gland and assessment of faecal testosterone in blackbuck. Keywords: Antelope cervicapra (L), Dominance hierarchy, Faecal testosterone, Indian antelope, Preorbital gland secretion, Scent marking behaviour Social dominance is a central issue in studies of marking from dominant males is far more effective in animal behaviour, and of great interest to behavioural promoting puberty acceleration in young female mice biologists. Dominance hierarchies often determine the than urine marking from subordinate males4. Since the first or the best access to food, social interaction, or social subordination suppresses gonadal function, mating within the animal groups1. The hierarchial emission of dominant male-specific chemical signals position of given individual can be influenced by its (pheromones) is under testosterone control. In age, body weight, aggressiveness, endocrine status addition, the subordinate males occupy less desirable i.e., androgen production, and/or scent habitats leading to lesser survival rate as well as lower marking/production2. Male dominance status is not reproductive potential4. simply a function of aggressiveness, but acquisition Scent marking by mammals is thought to be and maintenance of high dominance rank often important in the maintenance of territories and in involve frequent aggression moderated by conveying information among individuals in a rank testosterone which is considered as the quintessential order5. Scent marking frequency is correlated with 3 physiological moderator of such behaviour . Urine social dominance and the degree of aggressiveness in many mammalian species6. The scent marking ______ communication system of cervids, which is poorly *Correspondence: E-mail: [email protected]; [email protected] understood at present, has been the subject of several Supplementary data available online at NOPR along with the article studies in recent years. The compounds produced by 782 INDIAN J EXP BIOL, NOVEMBER 2018 various integumentary structures, such as the biological factors of dominant/subordinate animals interdigital, preorbital, forehead, and tarsal glands are have been subjected to a few studies, but there is no either carried on the body or deposited onto the information on the chemistry of the signals and the environment on olfactory signposts7. In antelopes, both role of faecal testosterone. We, therefore, investigated males and females possess preorbital organs which the frequency of preorbital gland marking and its have a glandular region in a pouch adjacent to the nasal chemical composition in dominant males before, (medial) corner of the eye8,9. There is no clear evidence during and after the dominance hierarchy period and of a role across species for the preorbital gland10 but the compared the data with bachelor and subadult males. most reported role of preorbital secretions is in This study is the first step in understanding the territorial marking11. It has been already proposed that potential role of preorbital scent gland marking in the the preorbital gland (POG) marking behaviour is dominance hierarchy of Indian blackbuck. commonly associated with reproductive activities, and is performed more frequently by harem males and may Materials and Methods serve to advertise the presence of a territory and to Study area attract males11. The study was conducted in the Conservation and In blackbucks [Antelope cervicapra (L)], only adult Breeding Centre of Arignar Anna Zoological Park males establish territories that are consistently (AAZP) (13°16’S and 79°54’E at an altitude of MSL+ occupied year-around12. The males perform scent 10-100 m), Vandalur, Chennai, Tamil Nadu, India. marking from their well developed POG, while in The Park covers an area of about 510 hectares. The females these glands are vestigial or functionless2,13. habitat of AAZP is considered a tropical evergreen In addition, the territorial male gland marking can scrub, a degraded forest mostly consisting of thorny serve as permanent threat signals and help to control bushes. The average annual rainfall is about 250 mm the subordinate behavioural activities such as scent and the temperature is about 26ºC with little marking behaviour, scent production, and patrolling seasonal variation. activity, in order to maintain the hierarchy14. We have earlier studied the urine scent marking behaviour and Study animals its chemical constituents of male blackbucks and At the beginning of the study period, the blackbuck found that the frequency of urine marking is enclosure housed a total population of 75 animals. All significantly higher in dominant males with specific- blackbucks were housed in an outdoor enclosure of urinary volatiles during the hierarchy period as about 3.5 acres (1.4 hectares) within a dry moat. The compared to that of bachelor and sub-adult males15. group was classified, according to Rajagopal & 2 As many as 28 major urinary compounds identified in Archunan , into 14 adult males, 20 adult females, 16 which three unique volatile compounds were present subadult males, 18 subadult females and 6 immatures. only in the urine of dominant males during the The dominance hierarchy system was studied in the dominance-hierarchy period. These specific-volatiles 14 adult males only (4-7 year old). Each individual may play a major role in the defense of territories was recognized by the difference in the shape of the against potential intruders and to advertise their horn and other morphological features (Table 1). agonistic dominance over other males15. Water was offered ad libitum, whereas food was Histomorphology of POG in territorial and non- served twice a day (10.30 a.m. and 4.00 p.m.). These territorial male Indian blackbuck indicates are exactly the same blackbuck individuals and same considerable histological changes in sebaceous and observation periods used in previous study and apocrine secretory glands, which would produce obtained data on scent marking and chemical 15 pheromonal substance through POG for olfactory constituents of urine . communication9. The preorbital gland-marking behaviour and the Social status of male blackbuck chemical components of gland secretion have been The nomenclature of male social status are: (1) harem identified for a few antelope (Bovidae) species masters (i.e. dominant males) who hold harems; including Neotragus moschatus16; Damalicus dorca (2) challengers (i.e. subdominant males) without their phillipsi17; Raphicerus campestris18 and Ourebia own harems, but they challenge the harem master and ourebi19. Further, the physiological, social and try to hold females; and (3) bachelor groups which stay RAJAGOPAL et al.: ESTABLISHMENT OF DOMINANCE HIERARCHY IN INDIAN BLACKBUCK 783 Table 1 — Antelope cervicapra L. special identification characters of individual adult male blackbucks Animal no. & No. of twists Identification characters (age)# in horn 1 (7 yrs old) 4* Narrow horns with pointed tips; a large size mole in the lower abdomen on left side 2 (5 yrs old) 3* Almost parallel
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