ELSEVIER

SEXUAL

BEHAVIOR
EJACULATORY

OF DONKEY JACKS: INFLUENCE

FREQUENCY
AND SEASON

OF

M.O. Gastal, M. H~~~Y,~A.R. Beker, E.L. Gastal and A. GonGalves3

Departamento de Zootecnia
Departamento de Veterintiia
Unpersidade
Federal de Viqosa, CEP 36570, Viqosa, MG, Brasil
Escola de Veterintia,
Universidade Federal de Minas Gerais
CP 567, CEP 30160, Belo Horizonte, MG, Brasil
Received for publication:
Accepted:

July

6,

Fehmr~y

1995

12, 1996

ABSTRACT
The courtship of 5 jacks was evaluated weekly for 12 mo. The characteristics of sexual
behavior were recorded before the collection of 2 ejaculates, at 4-h intervals, into an artificial vagina
in the presence of a female in either natural or induced estrus. The maximum time given to the jack
to perform the ejaculatory mount was 1h. If the jacks did not ejaculate, another attempt was made
the following week. At the time of collection, the male and female remained free in a paddock
(20 m2). The data presented in tliis study is based on results in which the first and second
ejaculates occurred successively. Statistical analysis was done by a 2 x 2 factorial design in
randomized blocks. The mean + SEM of sexual behavior characteristics for the first and second
ejaculate were, respectively: time until fist mount=0.7 f 0.2 and 0.9 f 0.2 min; time until first
erection=l1.9
f 1.1 and 11.6 + 1.1 min; time until ejaculation=15.0
+ 1.2 and 13.6 + 1.2 min:
frequency of partial exposure of penis=3.4 + 0.4 and 3.2 f. 0.4; frequency of total exposure of
penis=l.O + 0.2 and 1.1 & 0.2; frequency of flehmen responses=6.6 f 0.5 and 4.6 f 0.5; frequency
of erections=1.3 f 0.1 and 1.2 + 0.1; frequency of retreats away from the femaIe=l.8 + 0.2 and
1.0 f 0.2; frequency of mounts with erection but without ejaculation=O.3 + 0.1 and 0.1 f 0.1;
frequency of mounts without erection=2.0 + 0.1 and 1.2 * 0.1: and frequency of pelvic copulatory
movements=4.8 + 0.4 and 4.4 f 0.4. Individual differences were observed (PcO.05) for partial and
total exposure, flehmen responses, mounts without erection and pelvic copulatory movements.
The variables flehmen responses, retreats away from the female and morilts without erection
showed significant differences (PcO.05) between ejaculates. Seasonal effecp: on sexual behavior
characteristics were not found. However, a monthly effect was noted tar liehmen responses,
partial exposure of the penis, mounts without erection, retreats away from thr female and pelvic
copulatory movements when the 2 ejaculates were combined.
Key words: sexual behavior.

ejaculate , season, donkey jacks

Acknowledgments
This research was supported
by the Conselho National de Desenvolvimento Cientifico
e Tecnoldgico
(CNPq). The authors thank Dr. I. Sampaio and Dr. A.A. Oliveira fol
statistical assistance and Lisa Kulick for the preparation of figures.

Theriogenology
46.593-603,
0 1996 by Elsevier
Science

1996
Inc.

0093-691X/96/$15.00
PII 50093-691X(96)0021

1-5

594

Theriogenology
INTRODUCTION

Studies of the reproductive performance of donkeys are needed to increase their population
and thus avoid extinction of some of the breeds. These animals are very important to the economy
of certain countries since they adapt easily to arid regions, where they and their hybrids are used
frequently as a means of transportation,
Recent studies on the sexual behavior of animals have included not only the basic
physiologic mechanisms but also the suategies of mount used by the males in their natural
environment. Studies of sexual behavior include, in addition to the mount, all interactions between
males and females leading up to copulation (12).
Sexual behavior has been studied extensively in several domestic mammal species such as
bovine (7), porcine (ll), equine (2,4, 13, 16, 17,23, 24) and asinine (3,5, 9, 10, 14, 15, 18).
Research on jacks has verified that after the first contact with the female, the time necessary
for the male to carry out a mount and to ejaculate ranged from 6 to 32 min (1, 10, 14, 19, 20).
During the precopulatory and copulatory phases, jacks have been observed to exhibit behaviors
such as naso-nasal contact, flehmen responses, mounting without erection, partial and total
exposure of the penis, sniffing and biting specific areas of the female body, lip clapping (10, 14,
20). slow or quick reneats away from the female and vocalization (9,lO).
The objective of this study was to evaluate and quantify the sexual behavior characteristics
of jacks during courtship and to determine the seasonal effect on these characteristics over a 12-mo
period.
MATERIALS

AND METHODS

Animals
Five jacks (4 Nordestina and 1 of undetermined breed), 2 to 7 yr of age and weighing 120
to 200 kg, were used in the study. All animals were managed under the same conditions (5). The
males were submitted to natural luminosity and were fed with fresh sliced grass (Panicurn
n~~~irnunz)with variable nutritional quality, influenced by natural climatic conditions (latitude and
longitude of 195557S and 435632, respectively).
Sexual Behavior Evaluation
The sexual behavior of the donkeys was evaluated at the time of semen collection for
another study (6). For each behavior evaluation the male and the female were left free in a 20-m
paddock for a maximum of lh. After ejaculation the jack was removed from the paddock. Estrus
was induced in the jennies during the breeding season with Dinoprost Tromethaminea (5.0mg)
and during seasonal anestrus with Estradiol Cypionateb (4.0mg) due to the long experimental
period and the small number of available jennies. Two ejaculates, 4h apart, were collected
weekly with an artificial vagina for 12 mo. If the jacks did not ejaculate, another attempt was
%nalyse, Upjohn Phatmaceutical Products, Brazil.
bECP, Upjohn Phatmaceutical Products, Brazil.

595

Theriogenology

made the following week. Sexual behavior characteristics without an ejaculation were not
considered for statistical analysis. To record the behavior characteristics with a minimum of
human interference, the observer remained as far as possible from the animals during the
observation periods.
The following behaviors and parameters were recorded: 1) time until first mount- time
(minutes) after the fist contact with the female until the first mount with or without an erection;
2) time until first erection- time (minutes) after the first contact with the female until the first
erection with or without ejaculation; 3) time until ejaculation- time (minutes) after the first contact
with the female until the end of the ejaculation; 4) partial exposure of penis- number of partial
exposures of penis without exposure of the glans; 5) total exposure of penis- number of total
exposures of penis with total exposure of the glans, but without erection; 6) flehmen responsesnumber of upper lip reversions; 7) erections- number of penis erections including the ejaculation
erection; 8) retreating from the female- number of slow (walk) or fast (run) retreats away
from the jenny; 9) mount with erection- number of mounts with penis erection without
ejaculation; 10) mount without erection- number of mounts with partial or total exposure of penis;
and 11) pelvic copulatory movements- number of thrusts.
Statistical Analysis
A randomized block experimental design was used in the study. Analysis of variance was
based on a factorial scheme. The data were analyzed by the least squares method (8). The
mathematic model was
Yij k= P + Ai + E j + Mk

+ (AE)ij

+ (EM)jk

+ eiJk

= dependentvariable
p = mean
Ai = effect of animal(1,2,3,4,5)
E j = effect of ejaculate(1,2)
Mk = effect of groupof months(1,2,3,4, 5, 6)
(AE)ij = effect of interactionbetweenanimalandejaculate
(EM)jk = effect of interactionbetweenejaculateandmonth
eijk = randomerror
yijk

RESULTS
SexualBehaviorCharacteristics
The data were basedon the number of attemptsto collect ejaculateswhen the first and
secondsemencollectionoccurredon the sameday. The low numberof repetitiondid not allow
for the evaluationof interactionsof month by animalby ejaculate.Somecharacteristics
- partial
and total exposuresof the penis, flehmenresponses,erections,retreatingfrom the female, and
mountswith and without erections- were exhibitedregardlessof ejaculation;however, thesedata
were not usedfor this study. Other than the sexualbehaviorcharacteristicsanalyzed,behaviors
suchas sniffing and biting specific areasof the female body, lip clapping,vocalization, and
attemptsat masturbationwerealsoobserved.
The variables, partialandtotal exposuresof penis,flehmenresponses,mountswithout an
erectionand pelvic copulatory movements,showedindividual differences(P<O.O5).Moreover,

596

Theriogenology

differences(PcO.05) betweenfirst and secondejaculateswere found for flehmen responses,

retreatingfrom the female,andmountswithout erection(Tables1 and2).
SeasonalEffects
No seasonaleffect on sexualbehaviorcharacteristics
was found in this study (Figures 1
and 2). However, a monthly effect (PcO.05) was noted for somecharacteristicssuch as partial
exposureof the penis,flehmenresponses,
mountwithout erection,retreatingfrom the female,and
pelvic copulatorymovementswhenthe first andsecondejaculateswere groupedtogether(Figures
1 and2).
DISCUSSION
Generally,the first behaviorexhibitedby jacks in this study was vocalization,followed by
mountswith or without exposureof thepenis.Duringthis first contact,70.7% of the mountswith
exposuresof the penisoccurredwithout an erection,similar to the findingsof others (9, 14, 20).
In an earlierstudy(14) jacksliving without humaninterferencefrequentlyexhibited somemounts
with or without partial exposureof the penisat the beginningof courtship.Most of the sexual
behaviorsexhibited by thejacks during courtshipin this study have alsobeen observedby other
researchers(9. 10, 14, 15,20).
While the time prior to the fist mountwasbrief (Table 1) in our study, in an earlierreport
(10) in which thejenny wasrestrained,the time requiredwas 6.20 and4.64 min for the first and
secondejaculate,respectively.Apparently,thefact that the femalewas restrainedcontributedto an
increasein the time until the first mount in that study (10).
Our researchconfirmsthat the first erectiondoesnot alwaysresultin ejaculation.The mean
time until the first erection(11.8min) wassimilarto that found by Nishikawa(19), who reported
that approximately10.0mm werenecessary
for jacksto completean erectionafter the first contact
with a female.
The mean time until ejaculationdid not differ betweenanimalsor betweenejaculates,
unlike that in horses(23) in which thistimeperiodvariedbetweenanimalsand was higher for the
first ejaculate(7.37 min) thanfor the second(3.99 min). The resultsobtainedhere(Table 1) were
within the meanlimits cited by otherresearchers,
who alsodetectedthat the necessarytime for the
precopulatoryexcitation of jacks rangedfrom 10 to 32.6min (1,9, 10, 14, 18, 19, 20). However,
our resultsfor this variable tendedto be lower when comparedwith research(14) which used
jacks asa dummy (26 min), andwith anotherstudy (9) which evaluatedthe sexualbehavior of 2
jacks using approximately 20 jennies for each male (32.6 min). Comparing our results with
stallions(16, 23). it was obvious that jacks require a longer period (14.3 vs 3.7 min) of
precopulatory excitation. This variation could have been a difference between speciesand/or
management
factors.
Partialexposureof the penishasbeen reportedpreviously (10, 14), but only Henry et al.
(10) showedthe frequencyof thisevent (4.4) during the courtshipof jacks, which was similar to
our findings.The averageof total exposuresof the penis(Table 1) was low in our study, similar
to that of an earlier study with asses(10) in which when the peniswas completely exposedthis
wasgenerally followed by anerectionandejaculation.

1
2

1
2

1
2

10
10

07
07

08
08

06
06

07
07

0.4ti.5
0.5M.5

0.4fO.6
0.5kO.6

l.lkO.5
1.8f0.5

0.7M.6
1.3M.6

0.9~hO.6
0.4HJ.6

Time to first
mount
(minutes)
n mean+SEM

10
10

07
07

09
09

07
07

09
09

13.0?2.3
13.W.3

16.3U.8
10.M2.8

9.2B.4
8.8L2.4

12.5f2.7
14.452.7

8.4f2.4
12.lf2.4

Time to first
erection
(minutes)
n mean+SEM

10
10

07
07

09
09

07
07

09
09

15.2ti.4
14.7ti.4

20.7k2.9
11.7ti.9

12.3k2.5
10.2fl.5

16.1B.9
17.3fl.9

10.8ti.5
14.oti.5

Time to
ejaculation
(minutes)
n mearr&SEM

10
10

07
07

09
09

07
07

09
09

3.8?0.8
4.7ti.8

3.7+1 .o
2.0&1.0

3.7ti.9
3.3M.9

4.7fl.O
3.6fl.O

1.2M.9
2.3W.9

10
10

07
07

09
09

07
07

09
09

l.lM.4
2.0?0.4

0.4Hl.5
0.7M.5

0.7zhO.4
0.8H.4

2.2Ht.5
l.lM.5

0.4ti.4
0.6M.4

meadSEM

mean*SEM

No. of total
penis exposures

No. of partial
penis exposures

1
38
0.7kO.2
42 11.91H.l
42
15.M1.2
42
3.4M.4
42
1.W.2
2
38
0.9-10.2
42 11.6kl.l
42
13.6H.2
42
3.2M.4
42
l.lM.2
n = number of ejaculates.
a.b(P<O.OS)Values with different superscripts in the same column indicate signiftcant difference between ejaculates.

1
2

Total

1
2

Ejaculates

Jacks

42
42

10
10

07
07

09
09

07
07

09
09

6.6HJo.5a
4.6M.5 b

4.7&l .o
3.6H.O

3.4U.2
2.1f1.2

6.91.0
5.1H.O

14.1f1.2
8.6k1.2

4.0+_1.o
3.6kl.O

mean?SEM

No. of flehmen
responses

Table 1. Sexual behavior characteristics of jacks from the first contact with a female until ejaculation, based on weekly semen
collections over a 1Zmonth period.

1
2
1
2
1
2
1
2

10
10

07
07

09
09

07
07

09
09

1.4M.l
1.3HJ.l

1.2fo.2
l.lfl.2

1.4W.2
1.2ti.2

1.5M.2
1.5M.2

1.2ti.2
l.llfo.2

meadSEM

No. of erections

2.M0.6
0.6H.6

07
07

10
10

07
07
1.8fo.5
0.6ti.5

2.83.6
2.2ti.6

l.OItO.5
0.9W.5

1.1+0.5
0.8M.5

09
09

09
09

meadSEM

No. of retreats
from the female

10
10

07
07

09
09

07
07

09
09

0.4fO.l
O.lti.1

o.OdzO.2
O.WO.2

0.3kO.2
O.lzbO.2

0.7dzO.2
0.2ti.2

o.HO.2
O.lfi.2

meadSEM

No. of mounts
with an erection

10
10

07
07

09
09

07
07

09
09

3.1ti.3
2.01!0.3

1.5M.3
1.2ti.3

2.ofo.3
1.3ti.3

2.7M.3
l.lfl.3

0.9M.3
0.4M.3

mea&GEM

No. of mounts
without an erection

6.Ofl.7
5.6ti.7
4.OASl.6
3.8M.6

10
10

3.9zbO.7
2.8ti.7

3.OU.3
3.8f1.3

7.lz!O.6
5.9M.6

mean+SEM

07
07

07
07

02
02

08
08

No. of pelvic
movements

Total

42
1.8d10.2 42
0.3fo.I
42
2.M.l
34
4.8M.4
1
42
1.3f0.1
34
4.4W.4
2
42
1.2Ht.l
42
O.lfo.1
42
1.0k0.2b
42
1.2M.lb
n = number of ejaculates.
a,b(P~0.05) Values with different superscripts in the same column indicate significant difference between ejaculates.

1
2

Ejaculates

Jacks

Table 2. Sexual behavior characteristics of jacks from the first contact with a female until ejaculation, based on
weekly semen collections over a 12-month period.

599

Theriogenology

=
.P
El
a
z
I
5
G-l
8
f2
73
r
am
J-A

S-O

20 1

N-D
_

J-A

s-o

N-D

J-F

M-A

M-J

J-A

s-o

N-D

J-F

M-A

M-J

Month

J-F

M-A

M-J

N-D

J-A

:@:
bii
J-F

M-A

M-J

M-A

M-J

Month

J-A

s-o

N-D

J-F
Month

M-A

M-J

First ejaculate

J-A

J=number

q Second ejaculate ndumber

s-o

N-D

J-F
Month

of jacks
of ejaculates

Figure 1. Mean (ASEM) monthly of the sexualbehaviorcharacteristicsof jacks from the first
contactwith the femaleuntil the ejaculationundera weekly systemof semencollections
for 12 months.

600

J-A

s-o

N-D

J-F

M-A

M-J

J-A

N-D

s-o

M-A

M-J

M-A

M-J

Month

Month

0
1

J-F

J-A

S-O

N-D

J-F

Month

Month

First ejaculate

q Second ejaculate
J=number
knumber

of jacks
of ejaculates

Month

Figure 2. Mean (ASEM) monthly of the sexual behavior characteristics of jacks from the first
contact with the female until the ejaculation under a weekly system of semen collections
for 12 months.

Theriogenology

601

During courtship, the jacks showedan averageof 5.6 flehmen responses.A higher
frequency was observedfor the first ejaculatethan for the second(Table 1). Probably, the male
wasalreadystimulatedfor the secondejaculation.The averageof this variablewas similar to that
of anotherstudy on asses(6.6, 10). but tendedto be higherthat of horses(1.0, 16). Perhapsthe
longerperiodof precopulatoryexcitationrequiredby jackscontributedto thistime difference.
The mean number of times that the jacks showederections,including that before an
ejaculationwas 1.3. After eliminatingthe occasionsin which there were problemswith semen
collections,the causesfor the failure of ejaculationin somecasesremainedunclearin our study.
However, sometimesthe jacks made attemptsat masturbation(6.0 f 1.6/jack) without an
ejaculation.This behavior hasalso beenobservedin asses
(9) andin horses(16,25).
Slow (walk) or fast (run) retreatfrom the female is a behavior uniqueto jacks during
courtship. It appearsto be species-inherent
and is probably relevant to the sexual behavior of
donkeys.In our study, differences(P>O.O5)werenot observedbetweenjacks, but a larger number
of retreatswasdetectedfor the first ejaculatethanfor the second(Table 2). This behavior has
beencited by Henry et al. (IO), who detected1.17retreatsfrom thefemaleduring lh of courtship.
Mounts with erectionsbut without ejaculationoccurredat a lower rate (0.2) than in other
studies(1.39, 10). The primary causesassociated
with this event were refusal of the artificial
vagina by the male and rejectionof the male by the female.Mounts without an erectionwere
observedwith a meanfrequency of 1.6 times, and differencesbetweenjacks weredetected.The
first ejaculateshoweda higherfrequencyof mountswithout an erectionthanthe second(Table 2)
similarto the findings of others (10). In horses,a highernumberof mounts without an erection
wasalsoobservedbeforethe first ejaculation(23). Probablythe lower number of mountswithout
an erectionwas due to the animalshaving alreadyhadpreviouscontactwith the female.During
mounting,thejacks showedan averageof 4.6 pelvic copulatorymovements,similar to the results
(5.2) of Henry et al. (9). Individual differenceswereobserved(P<O.O5)but not betweenejaculates.
This resulttendedto belower thanthat reportedin stallions(7.0, 16).
A seasonal
effect wasnot detectedfor the time requiredfor the first mount, for the time
period prior to the first erection, or the time until ejaculation.Theseresultscontrast with other
studies(14, 19)in which a seasonal
effect on the libido of jacks was observed.According to these
researchers,the libido of jacks was more intensein spring-summerthan in autumn-winter.In
horses,Pickett andVoss(21) andPickett et al. (22,23) verified a shorterperiodof reactionduring
the physiologicalreproductiveseason.Seasonaleffect on sexualbehaviorcharacteristicswas not
evident in our study (Figures 1 and 2), sincethe monthly variation for some behaviors(partial
exposureof the penis,flehmenresponses,
mountswithout erection, retreatingfrom the female
andpelvic copulatorymovements)wereinconsistentduringcourtship.Maybe the lack of seasonal
effect on sexualbehaviorof jacks in our study could be associatedwith limited variation in the
weather(temperatureandphotoperiod)given the locationof this study.
In summary, we found that somecharacteristicsexhibited by jacks during courtship
differed betweenanimalsand that the first ejaculatewas a stimulantfactor for the second. A
seasonal
effect on sexualbehaviorwasnot evidenced,althoughmonthly variationswere observed
for somebehaviorsduringcourtship.

Theriogenology

602
REFERENCES

1. Bielanski W, Wierzbowski S. Some properties of the output of semen of jacks. Acta

Biol Cracov Ser Zoo1 1962;5: 117-124.
2. Bristol F. Breeding behavior of stallion at pasture with 20 mares in synchronised
oestrus. J Reprod Fertil 1982;32(Suppl):71-77.
3. Clayton HM, Lindsay FEF, Forbes AC, Hay LH. Some studies of comparative aspects
of sexual behavior in ponies and donkeys. App Anim Ethol 198 1;7: 169- 174.
4. Feist JD, Mccullough DR. Reproduction in feral horses. J Reprod Fertil 1975;23(Suppl): 1318.
5. Gastal MO. Estudo das caracteristicas seminais e do comportamento sexual de jumentos
MS Thesis, Belo Horizonte-MC, UFMG, 1991.
6. Gastal MO, Henry M, Beker AR, Gastal EL. Effect of ejaculation frequency and season on
donkey jack semen. Theriogenology 1995; in press.
7. Hafs HD, Hoyt RS, Batton RW. Libido, sperm characteristic, sperm output, and fertility of
mature dairy bulls ejaculated daily or weekly for thirty-two weeks. J Dairy Sci 1959;42:626636.
8. Harvey WR. Least-Squares and Maximum Likelihood Mixed Model. Ohio University,
1987. PC version (PC- 1).
9. Henry M, McDonnell SM, Lodi LD, Gastal EL. Pasture mating behaviour of donkeys
(Equus asinus) at natural and induced oestrus. J Reprod Fertil 1991;44(Suppl):77-86.
10. Henry M, Oliveira, MMF, Diaz AP, Gastal EL, Tolentino FT. Comportamento sexual de
jumentos no period0 de cortejo e ato sexual VII Brazil Cong Anim Reprod,
1987;7 1 abstr.
11. Hurtgen JP. Reproductive
Examination of the Boar. Hastings, NB: Society for
Theriogenology, 1984.
12. Keverne, EB. Reproductive
behaviour. In: Austin CR, Short RV (eds), Reproduction
in Mammals, vol. 4. Cambridge University Press, 1984; 133- 175.
13. Klingel H. Social organization and reproduction in equids. J Reprod Fertil 1975:23(Suppl):
7-11.
14. Kreuchauf A. Reproductive physiology in the jackass. Anim Res Devel 1984;20:51-78.
15. Lodi LD, Henry M, Paranhos da Costa MJR. Behavior of donkeys (Equus asinus) breeding
mares (Equus caballus) at pasture. VI Int Symp Equine Reprod, 1994;143-144.
16. McDonnell SM. Reproductive behavior of the stallion. Equine Pratt 1986;2:535-555.
17. McDonnell SM, Garcia MC, Kenney RM. Pharmacological manipulation of sexual
behaviour in stallion. J Reprod Fertil 1987;35(Suppl):45-49.
18. Morais RN. Contribuicao ao estudo da biologia reprodutiva de jumentos (Equus asinuS).
MS Thesis, Sao Paulo-SP. USP, 1990.
19. Nishikawa Y. Studies on Reproduction in Horses. Tokyo: Shiba Temuracho Minotoku,
1959.
20. Ostrowski
JEB, Cortesano JA. Production de mulares por inseminaccion artificial
Manejo de1 asno doador de semen. Rev Med Vet 1987;68:210-216.
21. Pickett BW, Voss, JL. Reproductive management of the stallion. Proc Am Assoc Equine
Pratt 1972;18:501-531.
22. Pickett BW, Faulkner LC, Seidel Jr GE, Bemdtson WE, Voss JL. Reproductive
J Anim Sci 1976;43:
1physiology of the stallion. 6. Seminalandbehavioralcharacteristics.
617-625.
23. Pickett BW, FaulknerLC, SutherlandTM. Effect of month and stallion on seminal
characteristicsandsexualbehavior.J Anim Sci 1970;31:713-728.

Theriogenology

603

24. Thompson Jr DL, Pickett BW, Berndtson WE, Voss JL, Nett TM. Reproductive physiology
of the stallion. 8. Artificial photoperiod, collection interval, seminal characteristics,
sexual behavior and concentrations of LH and testosterone in serum. J Anim Sci
1977:44:656-663.
25. Tischner M, Tomica E, Jezierski J. Age and seasonal effects on sexual behaviour of
stallions at rest. Anim Reprod Sci 1986;12:233-237.