AGRICULAGRICULAGRICULAGRICULAGRICULTURAL RESEARCH COMMUNICATURAL RESEARCH COMMUNICATURAL RESEARCH COMMUNICATURAL RESEARCH COMMUNICATURAL RESEARCH COMMUNICATION CENTRETION CENTRETION CENTRETION CENTRETION CENTRE

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SCENT GLSCENT GLSCENT GLSCENT GLSCENT GLAND AND ITS BEHAAND AND ITS BEHAAND AND ITS BEHAAND AND ITS BEHAAND AND ITS BEHAVIORAL FUNCTION IN SMALLVIORAL FUNCTION IN SMALLVIORAL FUNCTION IN SMALLVIORAL FUNCTION IN SMALLVIORAL FUNCTION IN SMALLMAMMALS– A REVIEWMAMMALS– A REVIEWMAMMALS– A REVIEWMAMMALS– A REVIEWMAMMALS– A REVIEW

Saroj BakshiSaroj BakshiSaroj BakshiSaroj BakshiSaroj Bakshi

Eco-toxicology Laboratory, Depertment of Zoology, Centre for Advanced Studies,University of Rajasthan, Jaipur - 302004, India

ABSTRACTABSTRACTABSTRACTABSTRACTABSTRACTVVVVVarious types of glands in small mammals werarious types of glands in small mammals werarious types of glands in small mammals werarious types of glands in small mammals werarious types of glands in small mammals were obsere obsere obsere obsere observed eg. lagmorphs (chingland, analved eg. lagmorphs (chingland, analved eg. lagmorphs (chingland, analved eg. lagmorphs (chingland, analved eg. lagmorphs (chingland, anal

gland, inguinal gland, hardenian, infraorbital, lacrymal gland), Columbian ground squirrels (scentgland, inguinal gland, hardenian, infraorbital, lacrymal gland), Columbian ground squirrels (scentgland, inguinal gland, hardenian, infraorbital, lacrymal gland), Columbian ground squirrels (scentgland, inguinal gland, hardenian, infraorbital, lacrymal gland), Columbian ground squirrels (scentgland, inguinal gland, hardenian, infraorbital, lacrymal gland), Columbian ground squirrels (scentgland at mouth and in anal area), Flying phalanger (frontal gland near ears and eye), Deer andgland at mouth and in anal area), Flying phalanger (frontal gland near ears and eye), Deer andgland at mouth and in anal area), Flying phalanger (frontal gland near ears and eye), Deer andgland at mouth and in anal area), Flying phalanger (frontal gland near ears and eye), Deer andgland at mouth and in anal area), Flying phalanger (frontal gland near ears and eye), Deer andBlack tailed deer (interdigital gland, metatarsal, tarsal gland) and Black tailed deer (interdigital gland, metatarsal, tarsal gland) and Black tailed deer (interdigital gland, metatarsal, tarsal gland) and Black tailed deer (interdigital gland, metatarsal, tarsal gland) and Black tailed deer (interdigital gland, metatarsal, tarsal gland) and Cricetus, Neotoma, PCricetus, Neotoma, PCricetus, Neotoma, PCricetus, Neotoma, PCricetus, Neotoma, Peromyscus,eromyscus,eromyscus,eromyscus,eromyscus,Meriones, TMeriones, TMeriones, TMeriones, TMeriones, Tatera, Rhombomys atera, Rhombomys atera, Rhombomys atera, Rhombomys atera, Rhombomys and and and and and RattusRattusRattusRattusRattus all species having midventral glandular organ. The all species having midventral glandular organ. The all species having midventral glandular organ. The all species having midventral glandular organ. The all species having midventral glandular organ. Thegland in males are significantly larger (P

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chemical signals originating in cutaneous scentglands.

The major function of scent marking isterritorial in nature as indicated by Hediger(1950). A number of other functions have beendesignated to scent marking, warn conspecificsaway from occupied territory, sex attractant orstimulant, labeling the home range for own useof orientation and maintaining a sense offamiliarity as an indicator of individual identityincluding information on sexual status, age,dominance and as indicator of population size(Johnson 1973). Extensive work on the scentmarking pattern and dominance hierarchy oflarge artiodactyls and musk shrew has beencarried out by Bala Krishnan and Alexander(1977), Pi l la i and Alexander (1987) andDominic (1987), (Table-1).

TTTTTypes of g lands in smal l mammals ypes of g lands in smal l mammals ypes of g lands in smal l mammals ypes of g lands in smal l mammals ypes of g lands in smal l mammals - Onespecies may possess varied assortment of glands.

Lagmorphs - Chin gland, anal gland, paired inguinalgland, hardenian, infra orbital, lacrymal gland.

Arctic and columbian ground squirrels - Scent glandat the corner of the mouth on the dorsal surface andin the anal area.

Flying phalanger - Major frontal gland and smallerglandular area near the ears and at the angle of theeye.

Cricetus - Mid ventral glandular organ

Neotoma - Mid ventral glandular organ

Peromyscus - Mid ventral glandular organ (Fig. 1,2)

Meriones - Mid ventral glandular organ (Fig. 3)

Rhombomys - Mid ventral glandular organ

Rattus - Twelve species of Rattus from the southpacific have been shown to possess ventral glandcomplex.

( R. exulana, R. annandalei, R. muelleri, R.Sabanus, R. rajah, R. surifer, R. cremoriventer,R . canus , R . edwards i , R . bowers i , R .jalorensis, R. meltada ) (Fig. 4)

Bat. Eonycteris spelaea – Para anal sebaceousglandular organs (Quay and miller – Schwarze 1970)

Microtene Rodents – Posterolateral sebaceousglandular region. Four standard sites of glandularenlargements and modifications caudal gland, rumpgland, hip gland and flank gland ( Quay 1968).

Deer, Odocoilrus virginianum – Cutaneous glandas lnterdigital gland, metatarsal gland, tarsal gland(Quay 1959) (Fig. 5)

Black-tailed deer Odocoileus hemionuscolumbianus - Integumentary glandular region aspreorbital, tarsal, metatarsal, interdigital, caudalregion (Quay 1970) (Fig. 6)

Caribou, Rangiefr- Glandular area as preorbitalgland, tarsal gland, interdigital gland (Quay 1955)(Fig. 7)

Kangaroo rat, Dipodomys – Glandular tissue (Quay1954)

Australian wild rabbit, Oryctolagus cuniculus –Submandibular cutaneous (Chin) gland, (Mykytowcz1966)

Antelopes, Oribi, Ourebia ourebia – Antorbital gland(Gosling 1972)

A spA spA spA spA specific midventral gland in Gerbils andecific midventral gland in Gerbils andecific midventral gland in Gerbils andecific midventral gland in Gerbils andecific midventral gland in Gerbils andRattus :Rattus :Rattus :Rattus :Rattus : The gland is situated on the midline of thebody, almost in the middle of the abdomen. Inappearance it is a small thickening of the skin twoto three times greater than neighbouring area andmany ducts empty into this region. The glandularfield is irregularly oval and elongated along themidline of the abdomen. The gland in males arelarger than those of females. The species of largeranimals with larger size of gland in comparision tosmaller size animals. The mid ventral gland caneasily be distinguished from the outside as the hairin the region is much shorter when pressed theglandular area a dense white secretion in somespecies or dirty yellow gland secretion with pungentsmell exudes from the ducts, musky odour more orless like that of kasturi of musk deer Moschusmoschiforus. The yellowish waxy exudation in the

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gland surface feels oily and has a distinctive muskyodour more or less like that of Kasturi of the muskdeer Moschus moschiforus. The gland is composedof complexus of enlarged sebaceous alveoli of typicalholocrine type, each with its ownduct, the gland unitsare separated from each other by thin layer ofconnective tissue which also support the walls ofthe ducts which are lined with stratified squamousepithelium.

The lower end of each duct opens intoseveral sebaceous alveoli. Holocrine secretioncollects in to common duct (Kumari et al. 1981).Mid ventral sebaceous complexes are common notonly within the genus Meriones but generally amongmembers of the family Cricetidae (Quay and Tomich1963; Thiessen and Yahr 1977). Nine of the 14recognized species of Meriones ( Nadler and Lay1967) are known to have midventral glands whichare longer and wider in males than in females(Sokolov and Skurat 1966; Thiessen and Yahr 1977).Both diurnal and nocturnal species have welldeveloped glands. The mid ventral gland of M.hurrianae is similar in many respects to that of M.unguiculatus. The histological structure of the glandspecialized sebaceous units associated with hairfollicles differ little between the two species (Glennand Gray 1965; Mitchell 1965) in M. unguiculatus.(Table 2)

The presence of the midventral gland enablesus to suggest a new systematic character for therepresentations of the subfamily Gerbillina.

Chemical nature of gland secretionChemical nature of gland secretionChemical nature of gland secretionChemical nature of gland secretionChemical nature of gland secretionA large number of macrocyclic chemical

substances have been identified as being producedby mammals exudative gland and are observed tobe pheromonal in function. The first substancesidentified were the musks produced by para analpouches and preputial glands of musk deer (Moschusmoschiferus). The tarsal scent glands of black taileddeer (Odocoileus hemionus) secrete a number ofcompounds including one chemically identified asa cis-4-hydroxy-doctec-6-enoic acid lactone (Muller-

Fig.2 :Fig.2 :Fig.2 :Fig.2 :Fig.2 : Typical size and location of the ventral scent glandfor a male (left), lactating female (middle), andnonpregnant female (right) in Mongolian gerbilMeriones unguiculatus (Kittrell, Gregg andThiessen 1982)

Schwarze 1971). Goodrich and Mykytowcz (1972)examined the compounds produced by various skinglands in the rabbit (Oryctologus cuniculus). Thechin glands produce primarily proteins and

Fig.1Fig.1Fig.1Fig.1Fig.1 : Mature male (A) and female (B) Peromyscusmaniculatus bairdii midventral sebaceous area(MSA), (Doty and Kart, 1972) (Left)

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carbohydrates and the concentration of them is farhigher in males than it is in females. The secretionof inguinal gland is composed primarily of lipids withmuch monoglyceride, diglyceride and triglyceridematerial. The secretion from the flank organs of watervole (Arvicola terrestris) is composed of complexmixture of at least 50 main components. Thecomponents are homologous series of isomers of longchain alcohols (Stoddart 1976). A highly volatileconstituent of ventral gland secretion of theMangolian gerbil (Meriones unguiculatus) isphenylacetic acid (Thiessen et al. 1974). Singer etal., (1976) have reported dimethyl disulfide isolatedfrom vaginal secretion of golden hamster(Mesocricetus auratus) The male preputial glandapparently secrets a signalling pheromonal (Bronsonand Caroom 1971) which acts olfactorily, attractingfemales under the restricted conditions of laboratoryshowed that the preputial gland factor was attractiveto female mice, whether it was contained in urine inthe form of a saline homogenate or occurred as alipid extract. Lederer (1950) pointed out thatcastoreum from the beaver (Castor fiber L.) containedno less than 45 different substances. Stevens andErickson (1942) identified the main components ofthe musk from the musk rat (Ondatra zibethicus) ascyclopentadecanol and cycloheptadecnol and theircorresponding ketones.

HistologyHistologyHistologyHistologyHistology, se, se, se, se, sex and age rx and age rx and age rx and age rx and age relationshipelationshipelationshipelationshipelationshipThe gland is significantly (P

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Fig.5 :Fig.5 :Fig.5 :Fig.5 :Fig.5 : The location of the glandular areas in Deer, Odocoileus virginianus diagrammed here; from left to right:forefoot interdigital gland, hind foot interdigital gland, metatarsal gland. (Quay 1959)

TTTTTable 1 : able 1 : able 1 : able 1 : able 1 : Responses to scent marks in mammalsResponses to scent marks Source

DefDefDefDefDefense against predatorsense against predatorsense against predatorsense against predatorsense against predators Stoddart (1976)

Social communicationSocial communicationSocial communicationSocial communicationSocial communication Stoddart (1976)

1) Population membership

2) Family membership

3) Sex

4) Sexual condition

5) Dominance status

Own markOwn markOwn markOwn markOwn mark

1) Increased confidence or reduced anxiety Ewer (1968)

2) In social species may also increased friendliness or decreased aggression

FFFFForororororeign markeign markeign markeign markeign mark Ewer (1968)

1) Increased confidence readiness to flee

2) Increased aggression, readiness to fight

3) Where there is recognition of individual marks, may reduce the effect of familiar marks

4) Where the setter of the mark and the response are of opposite sex, a foreign mark

may have a positive effect, causing sexual arousal and producing appetitive behavior for mating

5) In social species the usual negative effect may be combined with, or replaced by a positive effect

so that even the marks of members of a foreign group may not be avoided but act as pathway signals.

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Meriones unguiculatus, Rhombomys opimus(Sokolov and Skurat 1966). Presence of gland onlyin male Rattus exulans (Quay and Tomich1963), Rattus melteda (Prakash and Kumari1979) (Table 3).

Histopathologically these gland is composedof complexes of enlarged sebaceous alveoli of typicalholocrine type, each with its own duct. The glandunits are separated from each other by thin layer ofconnective tissue which also supports the walls ofthe ducts which are lined with stratified squamousepithelium. The lower end of each duct open intoseveral sebaceous alveoli. The basal alveolar cellsare small but as they develop they become

progressively enlarged, their nuclei shrink anddisappear and cells break down into fatty detrituswithin the luman of the holocrine secretion collectsinto a common duct through which the hair alsopass. The hairs flattened curved surface which isconsidered help for the movement of sebum out ofcells (Kumari et al. 1981; Sokolov and Skurat 1966;Doty and Kart 1972).

Age and gland size correlated (P

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of male and female desert gerbil with their bodyweight . The g land s i ze d id not d i f fe rsignificantly between subadults of two sexes butin adult the heavier body weight classes maledesert gerbil had larger glands and with largerareas than females more so in 81-100g bodyweight class corresponding to 8-12 months ofage (Table 4&5). It is evident that gland sizeincreased markedly in both males and femalesupto 80g body weight and there after increasedsignificantly only in male. When gland lengthand width were regressed separately againstbody weights for each sex and the regressioncoefficients for each measurement comparedbetween sexes, males differed significantly(P

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TTTTTable 2 : able 2 : able 2 : able 2 : able 2 : Dimension of mid ventral gland among Desert Rodents.

Species

Sex

Gland Length (mm)

Gland Width (mm)

GlandArea

(mm2

1

2 3 4 5

Family – Muridea Sub - Family – Gerbillinae

Meriones unguiculatus

Meriones erythrourus

Meriones tamariscinus

Meriones tristrami

Meriones persicus

Meriones meridianus

Meriones hurrianae

Tatera indica indica

Rhombomys opimus

Sub – Family – Murinae Rattus exulans

Rattus meltada

Family – Cricetinae

Peromyscus maniculatus artemisiae

P b i dii

Male Female Male

Female Male

Female Male

Female Male

Female Male

Female Male

Female Male

Female Male

Female

Male Female Male

Female

M l

23.4 16.1 29.0 21.0 29.5 15.5 25.0 21.0 19.0 13.0 13.0 7.0

21.17 14.86 17.38

- 22.00 15.00

43.00 -

22.17 -

17 63

5.0 3.2 7.0 5.0 6.5 3.5 6.0 6.0 6.0 3.0 5.0 2.0 4.87 2.72 3.28

- 7.00 4.00

4.7 -

5.96 -

1 88

115.051.52203.0105.0191.754.25150.0126.0114.039.065.0014.00109.341.5958.59

- 154.060.00

172.0-

160.8-

33 14

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the number of individuals per unit area suggesting asimilar decrease in the rate of scent marking(Table 7). Normal scent marking in cages undervarying densities also reduced to about 66.6 %indensity of three and 33.3 % in density of 6.9 and12 percage (90 x 30 x 30 cm.).

Scent marking and social hierarchyScent marking and social hierarchyScent marking and social hierarchyScent marking and social hierarchyScent marking and social hierarchyAnother factor which has come to light for

explaning this differences in frequency of scentmarking is their dominance behaviour. As therodents are grouped in laboratory cages or in therattery a male or a female acquires dominance statusthrough fighting, chasing and scent marking at ahigher rate. This finding was further investigated byplanning a series of experiments in plus mazes, glass

cages and the rattery. In the rattery a group of 3male and 6 female M. hurrianae was released. All ofthem were marked by toe-clipping and dye markedto facilitate identification of each desert gerbil. Soonthe release they started digging activity and in about15 days time a cohesive social group was formed.

The dominant male as well as female wereidentifiable became of the number of times othersstayed away from them and on the basis of theirchasing activity, which is an important componentof their aggressive behaviour as compared to fighting(Table 8).

Male dominance scent marking has beenobserverd in Merionus tristami (Thiessen et al. 1973),M. ungiculatus (Thiessen and Yahr 1977), M. libycus

TTTTTable 3 : able 3 : able 3 : able 3 : able 3 : Body weight classes and gland size of male M. meltada (after Kumari $ Prakash 1983)Body weight classes (g) Body weight

(g) Gland length

(mm) Gland width

(mm) Gland area

(mm2)

30-50 51-70 71-90 91-110

41.25±3.28 62.82±1.09 79.06±1.2 97.50±1.0

21.9±2.8* 24.6±1.0* 29.5±3.1NS 30.7±4.0

5.95±0.47** 6.05±0.26*

7.06±0.44NS 7.50±0.087

139.0±23.4 155.3±12.2** 223.40±16.42* 239.50±57.9

Body weight classes (g) Body weight (g)

Gland length (mm)

Gland width (mm)

Gland area (mm2)

40-60 61-80 81-100 101-120 121-140 141-160 161-180

49.90±2.6 69.94±1.64 90.83±1.5

111.79±5.79 132.86±1.86 148.29±1.97 168.16±1.29

9.33±0.95* 11.64±1.44* 16.11±1.38* 18.94±1.58NS 19.41±1.09NS 20.54±2.54NS 22.09±1.41

1.83±0.24NS 2.42±0.35NS 3.07±0.34* 3.33±0.34* 3.76±0.34NS 3.75±0.04NS 4.08±0.36

17.83±0.86** 28.96±5.20** 50.03±4.35* 57.65±4.51** 73.75±8.47* 76.95±10.04* 90.27±9.37

T test; level of significance between next body weight class *P

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and M. crassus (Daly 1977). Male dominance alsoreported in rabbit Oryctologus cunniculus(Mykytowycz 1965), black tailed deer Odocoileusherrionus columbianus (Muller Velten 1966) and inmarmot Marmota maromota (Munch1958).

Adaptive value of scent marking in regulatingAdaptive value of scent marking in regulatingAdaptive value of scent marking in regulatingAdaptive value of scent marking in regulatingAdaptive value of scent marking in regulatingpopulation mechanismpopulation mechanismpopulation mechanismpopulation mechanismpopulation mechanism

The more severe magnitude of dominancein female M. hurrianae as demonstrated by sebummarking is visualized in the context of evolutionof sociability and adaptive value of self regulatingpopulation mechanism (Wynne Edwards 1962;Eisenberg 1967) where their numbers are kept

below a maximum level by restricting mating toonly through social interactions the dominantmale to mate with other females in the socialgroup. It is also possible that the dominant femaledesert gerbil familiarizes the alpha male with itsown odour thus obliterating other females fromchance of mating (Kumari and Prakash 1981b).A number of view’s have been advanced byvarious workers regarding the behaviouraladjustments with in a species for self regulatorymechanisms of population control. Chitty (1960)suggested a genotypic hypothesis but it was notcorroborated by other ethologist Christian andDavis (1964) support phenotypic hypothesis of

Family

Species Source of Marking

Aplodontoidea Sciuroidea

Castoroidea Muroidea

Gliroidea Hystricoidea

Erethizontoidea Cavoidea

Octodontoidea

Aplodontia rufa Sciurus Xerus

Marmota

Citellus Castor

Circetus Mesocricetus

Arvicola

Ondatra Rattus

Mus

Notomys

Cricetomys

Glis Atherurus Erethizon

Cavia

Dasyprocta

Myoprocta

Thryonomys

Urine Urine Cheek glands: rubbed on objects Cheek glands: rubbed on objects Anal glands: anal drag Cheek glands: rubbed on objects Anal gland: rubbed on objects Lateral gland: transferred to ground by rubbing foot Lateral glands: rubbing of foot on gland followed by stamping Anal gland : anal drag Urine: drip trail and may mark defeated rival mid ventral gland in some species Urine: drip trail and may mark defeated rival foot gland Ventral gland on chin or chest: usually larger in male then female Urine : drip trail – only when disturbed Faces : hand stand posture seen occasionally when disturbed and then only in male Salive licking combined with rubbing Cheeks on Prominent objects. Cheek glands Commoner in male then female: male may mark female during courtship Anal: anal drag Anal gland : anal drag: seen only in male Cheek glands : rubbed on objects Anal gland: anal drag may be combined with scratching commoner in male then in female Anal gland: anal drag may be combined with scratching commoner in male then in female Anal gland: anal drag may be combined with scratching commoner in male then in female Cheek gland: rubbed on objects Anal gland: anal drag may be combine anal marking and scratching the ground offer a victory (seen only in male)

TTTTTable 6 :able 6 :able 6 :able 6 :able 6 : Marking method in Rodents (After Ewer 1968).

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self regulation by a behavioural endocrine feedback system elicitated by ‘Social pressureSocial pressureSocial pressureSocial pressureSocial pressure’ andoperating through the pituitary adrenocorticalaxis.

Scent marking and homoestasisScent marking and homoestasisScent marking and homoestasisScent marking and homoestasisScent marking and homoestasisThe diurnal desert gerbil, M. hurrianae

lives in complicated burrow system in whichalmost a constant and comfortable temperatureis maintained (Prakash 1981). After exposure tothe hot outer environment during the day therodent develops hyperthermia and it affloads theexcessive heat load into the cooler environmentof the burrow by intermittently visiting it (Prakash1986). In this context the glandular region couldhas a “ heat windowheat windowheat windowheat windowheat window” allowing an exchange ofheat with the surrounding environment whensebum is exuded out (Kittrell et al. 1982). Thescent mark ing process , thus ass i s t shomeostasis in temperature regulation besideschemocommunication. This function of the

scent marking gland may explain the reductionof scent marking frequency when they are in cagesand are grouped together since due to huddlingthe necessity for homeostasis also declines.

Scent marking response to conspecific odoursScent marking response to conspecific odoursScent marking response to conspecific odoursScent marking response to conspecific odoursScent marking response to conspecific odoursThe rodents were given choice to react to

glass slide smeared with their own sebum odour ofstrange male and strange female. A plain glass slidewas placed on the opposite side of the cage or in adifferent arm of the plus maze as control. Surprisinglyno sex attraction was observed as males preferredthe unisex odour as adjudged by their own scentmarking frequency, number of visits and duration ofstay. Only in the absence of same sex sebum odour,they preferred odour of opposite sex as comparedto behavioral acts performed near the blank slide.Both sexes of M. hurrianae even preformed unisexsebum odour of another gerbil. This behaviour ofM. tristami (Thiessen et al. 1973). Under similarchoice test combinations the Mongolian gerbils show

TTTTTable 7 : able 7 : able 7 : able 7 : able 7 : Frequency of sebum marking per 15 min, average of three replication (Mean±SE) (Kumari $ Prakash 1984)

Scent marking frDensity in cage 90x30x30 cm M. hurrianae

Individual Group of 3 Group of 6

Group of 9/12

9.75±4.98 4.41±2.76* 2.41±1.17* 1.55±0.78NS

Level of significance between the group and the one above (Student’s test) **P

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a significantly preferential response toward odourderived from other males as compared towardfemale sebum odour. However the females of M.hurrianae show a significant preference for femaleodour where as females of M. unguiculatus and M.tristrami do not indicate significant preference forthe odour of either similar or opposite sex (Thiessenet al. 1970,1973). Daly (1977) suggested that in M.unguiculatus marking has a familiarizing functionassuming access to females during proestrous andhence giving the male a chance to be first on thescene during oestrus.

In absence of clear sex attractant functionof sebum odour in the three rodent species it hasbeen explained that an animal own scent might actto ‘increase its confidanceincrease its confidanceincrease its confidanceincrease its confidanceincrease its confidance’ in the environment(Ewer 1968) where as Eibl-Eibesfeldt (1953) andMykytowycz (1968) conjecture that scent marksprovide ‘homelinesshomelinesshomelinesshomelinesshomeliness’ to the animals. It has beenobserved that in the field as well as in large ratteryM. hurrianae and T. indica ventral mark the burrowopenings quite often especially before entering theburrow. These rodents also scent mark foodcontainers, the grass clumps in their home rangeand any new object whether it is a stone or a wooden

peg. From these observations it appears that thefunction of scent mark is more of a ‘familiarizationfamiliarizationfamiliarizationfamiliarizationfamiliarization’’’’’nature or to signal home to the marking rodent orthat of labeling the habitat for an animals ownuse inorientation (Johnson 1973). Since food is scenty inthe desert and its availability is a prime for theirsurvival there is a keen competition for it. The scentmarking of grass clumps in the field and foodcontainers in the rattery by the two gerbils suggestsa ‘food rfood rfood rfood rfood reseresereseresereservationvationvationvationvation’’’’’ function of the sebum odour(Kumari and Prakash 1981a & b) (Table 9).

Sebum marking and ReproductionSebum marking and ReproductionSebum marking and ReproductionSebum marking and ReproductionSebum marking and ReproductionThe female desert gerbils when in oestrus

increase the scent marking frequency and it isperceived by male gerbils almost by doubling theirmarking activity (Kumari and Prakash 1981,1984a).The average period of oestrous cycle in M. hurrianaeis of 6.2 days (Ghosh 1968) which was determinedby examining vaginal smears followed (Allen1922).The scent marking behaviour of female desert gerbilswas observed in glass cage (92X31X31cm.) for 15minute daily at 1100 hours. The introduced femalescent marked in the clean cage for observing theresponse of male toward female odours of dioestrusfemales.

Intraspecific communication Interspecific communication

Individual appraisal Group membership appraisal

Age appraisal Social status appraisal

Sex appraisal Reproductive stage indication

Trail marking Territory marking

Identification with home range Warning Defense Alarm

Submission Attention – seeking

Greeting Encouraging approach

Distress- signaling Pain indication Gregariousness

Individual Species membership

Prey Predator Warning Defense

TTTTTable 9 : able 9 : able 9 : able 9 : able 9 : Types of messages which could be conveyed by mammals by means of olfaction.(Schultze – Westrum, 1965)

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The frequency of sebum marking by thefemale M. hurrianae increased significantly(P

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throughout the year irrespective of the breedingseason as in Siberian hamster (Vorontosovand Gurtovoi 1959), black-tailed deer (Muller-Schwarze 1971), kangaroo rat (Quay 1953),Mangolian gerbil (Mitchall 1967), White rat(S tandy and Powel l 1941) and severa lar tiodactyles (Von-Schumacher 1936; Moy1969). The results of our studies provide rather

inconclusive evidence about the relationshipbetween breed ing per formance and thefluctuations in the gland size of these threerodent species (Kumari and Prakash 1983).

ACKNOWLEGEMENTACKNOWLEGEMENTACKNOWLEGEMENTACKNOWLEGEMENTACKNOWLEGEMENTI am greatly thankful to Miss Princy Shakeet

for collecting and compiling the data for themanuscript.

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