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Showing posts with label vagina. Show all posts
Showing posts with label vagina. Show all posts

Saturday, November 30, 2019

A brief look at the family Leporidae (hares and rabbits)

Keywords: Leporidae, Oryctolagus, Lepus, Sylvilagus, anatomy, physiology, reproduction, uterus, ovaries, tract, vagina, rabbits, hares

A fascinating group of mammals, some familiar to veterinary and animal science, others not. Note that  rabbits and hares belong to the order Lagomorpha, of which there are two families: Leporidae and Ochotonidae (pikas, mouse hares, or conies)

As mentioned in the LORI entry on semen collection in rabbits, rabbit farming is underdeveloped in North America compared with Europe, Asia and South America. Therefore rabbits are superficially addressed in most animal science and veterinary curricula in North America. However, rabbits deserve attention as important sources of meat and fiber and their unique value as subjects for scientific study (See: Foote and Carney, 2000). 

Vocabulary:

Lagamorph: A member of the order Lagamorpha
Leporid: A member of the family Leporidae, comprising the rabbits and hares.
Kindling: The name given to parturition in rabbits and hares.
Kits: Perinatal rabbits and hares (the term "pup" is reserved for rats, mice and dogs).
Doe: Adult female Leporid
Buck: Adult male Leporid.
Nests and burrows: Hares (always wild) live in nests, above ground. Wild rabbits such as Cottontails live in burrows.

Introductory notes: Rabbits and hares are all members of the family Leporidae.  This family includes the domestic rabbits (Oryctolagus cuniculus), hares (many Lepus spp) and brush rabbits (many Sylvilagus spp). They share similar reproductive characteristics but as donkeys are to horses, they are also distinctively different. For example, hares and brush rabbits have 48 chromosome and precocious neonates with open eyes, able to ambulate efficiently. By contrast, rabbits have 44 chromosomes and altricial young, blind, hairless and unable to ambulate. Their eyes open at about 10 days postpartum.

In the course of this entry, the author will entertain other differences between hares and rabbits. 

Figure 1. The reproductive tract of a Snowshoe hare (Lepus Americanus) that was approximately 14 days pregnant. The ~20 day old fetus at left is that of a domestic rabbit (Oryctolagus cuniculus).  The 14 day pregnancy shows the bulbous embryo-placental units typical of rabbits and hares. Note too, that the ovaries are not covered by ovarian bursae. Corpora lutea of pregnancy are visible within the ovaries. Finally, note the double cervix (inset at right); typical for all Leporids (as well as rats and mice). The vagina (not shown) is about 10 to 12 cm long. Image size: 1344 x 831 px

A crash course on rabbit and hare reproduction: 
1. Puberty occurs between 4 and 7 months of age; later in large rabbit breeds such as the Flemish White. 

2. Although commercial rabbits breed throughout the year, hares and wild rabbits (Sylvilagus spp; Cottontailsare like cats. i.e. they respond to increasing day length and breed mostly during spring and early summer. 

3. Like cats, rabbits and hares do not have discrete estrous cycles. Instead, as reflected by vaginal cytology, follicle waves mature every 4 to 5 days (~ 7 days in hares). 

4. Although spontaneous ovulation can occur, ovulation is generally induced by copulation (mounting in the absence of intromission is also thought to be important). Ovulation occurs about 10 hours post copulation. GnRH treatment is essential for satisfactory pregnancy rates when AI is used.

5. Pregnancy diagnosis is accurate at 10 to 14 days post-breeding using transabdominal palpation. Optimal first service conception rates should be greater than 70%  i.e. 70% of does should be pregnant after a single breeding.

6. Gestation is longer in hares than rabbits; about 42 days vs. 30 days. This explains why hares have precocious young and rabbits, altricial young. 

7. Parturition occurs in the morning. Anterior, posterior and even breech presentations are normal. Usually kindling is complete in less than 30 minutes but occasionally, kits (like kittens) may be born hours or even one or two days apart. Kits are usually born separated from their umbilical cords but maternal biting of cords has been described.

8. Cannibalism (infanticide) is fairly common in young rabbits, especially under stressful conditions.

9. Rabbits have an average of six to 10 kits per litter. Does have 8 to 10 mammary glands and nurse their kits once per day.

10. Pseudopregnancy occurs after sterile mating in both hares and rabbits but does not normally occur in the absence of mating as in dogs and cats. Like pregnancy itself, pseudopregnancy is slightly longer in hares than rabbits i.e. 24 days vs. 17 days. As is sometimes seen in bitches, nesting behavior will occur during pseudopregnancy. In the case of Leporids this involves pulling of abdominal hairs and gathering of straw etc. 

11. Pregnancies with one or two fetuses may be significantly longer than otherwise (as is the situation with cats and dogs). One can usually diagnose retained fetuses by palpation. Cesarean section may be required in such cases.

12. Rabbits can have fertile copulation soon after birth and become pregnant in under a week after parturition (kindling ). This is possible because rabbits have active ovaries at the time of kindling and uterine involution is largely complete within 48 hours of parturition. Consider that other domestic animals have ovaries devoid of significant follicle growth at parturition.

13. Conception soon after parturition is remarkable in light of the intimate placentation in Leporids. It is hemochorial i.e. more intimate than any other domestic animal (see figure 2).
In commercial operations, does are re-bred between 2 and 6 weeks after kindling. This will provide 4 to 6 litters per year.

14. Weaning in commercial operations usually occurs at approximately 6 weeks, when the doe has already been pregnant for some time.

15. Rabbits are usually processed for meat production at 3 to 4 months of age and breeding stock  rabbits are seldom kept for longer than 3 to 4 years in commercial units.



Figure 2. Placentation typical of a rabbit or hare; a modified amalgum from several sources. Placentation is bi-discoidal then discoidal and intimate, developing mesometrially (although anti-mesometrial in early gestation). Placentation is hemochorial (also referred to as hemobichorial) with giant cells forming in the endometrium, similar to equine eCG-producing cells. Note the large exocelom, similar to that in carnivores. Image size: 896 x 733 px

Selected references:

Benirschke. K. Comparative placentation. http://placentation.ucsd.edu/rabbits.htm

Chavatte-Palmer P. et al. 2008. In utero characterization of fetal growth by ultrasound scanning in the rabbit. Theriogenology. y 69: 859–869

Dickie, E. 2011.Dystocia in a rabbit (Oryctolagus cuniculus). Can. Vet. J. 52: 80-83.

Boumahdi, Z. et al. 20009  Behavior at birth and anatomo-histological changes studies of uteri and ovaries in the post partum phase in rabbits. European J. of Scientific Res. 34: 474-484

Foote, R.H. and Carney E.W. 2000.The rabbit as a model for reproductive and developmental toxicity studies 

Foxcraft, G.R. and Hasnain, H. 1973. Effects of suckling and time to mating after parturition on reproduction in the domestic rabbit. J. Reprod. Fert. 33:367-377

International rabbit reproduction group. 2005.Guideline for the handling of rabbit bucks and semen. World rabbit.Sci. 13:71-91

Roellig, K. et al 2010. Superconception in mammalian pregnancy can be detected and increases reproductive output per breeding season. Nat Commun. 2010 Sep; 1(6): 1–7.

Torres, S. et al. 1977. Fertility factors in lactating rabbits mated 24 hours and 25 days after parturition. Ann. biol. anim. Biochem. Biophysics. 17:63-69

Weisbroth et al Eds. 2013. The Biology of the Laboratory Rabbit. ISBN 1483270319

Monday, March 20, 2017

Rabbit semen collection and use.

Rabbit semen collection and use.

Keywords: rabbit, semen, collection, vagina, artificial, lagomorph

Compared with Europe, Asia and South America, rabbit farming is underdeveloped in North America. Therefore, it is only superficially addressed in most of our animal science and veterinary curricula. Nevertheless, rabbits are important as livestock; farmed for meat and of course, angora fiber. They also have unique value for humane scientific study (See: Foote and Carney, 2000). 

On one occasion, guided by ignorance, the author built this surprisingly effective artificial vagina (AV) to collect rabbit semen. It was based on a bovine AV but had a liner constructed from the finger of a surgical glove. A small hole on the side, covered by another piece of the glove was created to admit warm water and air as required. As can be seen, the liner terminated in a small collection vessel, cut from the conical tip of  plastic test tube. 


Image size: 1151 x 1563 px

Using a receptive doe, the process of semen collection was simple. Semen collection can be further simplified by training the buck to mount the skin of a doe, covering the hand that holds the AV. Copulation is rapid, resembling most closely, that seen in ruminants.

In the majority (~70%) of semen samples, a small plug of clear mucus will be found in the AV as shown below. This is a product of the vesicular glands. In sequential samples collected on the same day, the presence of a mucous plug is less likely (~5%). It was once surmised that the plug formed by this gel was essential to prevent loss of semen and produce normal conception rates. However it is now clear that conception rates can be excellent in the absence of any gel plug. Indeed, extended semen contains virtually no gel and conception rates using A.I. can be excellent. Therefore the presence of gel in rabbit ejaculates remains an enigma, rather like the gel in a stallion's ejaculate.


Image size: 1625 x 1099 px

To the author's eye, rabbit semen bears a striking resemblance to that of a bull; its motility and morphology being similar. Concentration is lower that of a bull's ejaculate however (~500 million per ml vs. 1000 million per ml). Nevertheless, wave motion (a function of both motility and concentration) can often be seen in rabbit ejaculates. 

Puberty in males is reached at six to seven months of age but peak sperm production may not be reached until 2 years of age or more.  At maturity, the volume of an ejaculate varies from 0.5 to 2 ml (occasionally more) but there is some variation at both upper and lower limits, partly due to significant volumes remaining on the liner of the AV.  For that reason, the AV is sometimes flushed after collection to retrieve more of the ejaculate.

Although sperm production increases under the effect of increasing day length in wild lagamorphs, this effect is not generally noticeable in domestic rabbits.

Although AVs are commercially available, readers may be interested to read about an alternative AV (Bredderman, 1964) that can be built for small scale breeding. Semen extenders and insemination pipettes are widely available for commercial rabbit production. Sources are readily available on the Internet.

Handling and insemination

Although acceptable fertility has been reported with doses as low as 0.5 million sperm, common practice dictates that between 20 and 40 million sperm are used per insemination, aiming for no more than 0.5 ml in volume to reduce sperm loss after insemination. A single ejaculate therefore, may provide between 20 and 50 insemination doses. 

For commercial use, semen from several males is pooled before insemination. The insemination pipette is bent dorsally to accommodate the ventrally-directed vagina and semen is simply deposited in the cranial vagina. No attempt is made to inseminate intra-cervically in either of the two cervixes. 

Frozen-thawed semen is seldom used in commercial rabbit production. Instead, ejaculates are diluted in commercial diluents, cooled to between 18 and 20 deg.C and used over a 24 to 48 hour period. 

To ensure that ovulation occurs (bearing in mind that it is predominantly mounting and not vaginal stimulation that induces ovulation in rabbits; Cf. cats) it is common to inject small intramuscular doses of GnRH analogs into the doe at the time of insemination. Because GnRH and its analogs are small molecules and readily absorbed through mucous membranes, some success has been achieved by adding GnRH analogs (usually buserelin) to insemination doses to induce ovulation. Without GnRH stimulation, the number of kits per litter is not commercially viable.

Selected references:

Alvarino, J.M.R. 2000. Reproductive performance of male rabbits World Rabbit Science Association, Volume 8, supplement 1. pp.13-35

Amann, R.P. 1966. Effect of ejaculation frequency and breed on semen characteristics and sperm output of rabbits. J. Reprod Fert. 11:291-293

Bredderman, P.J. 1964. An improved artificial vagina for collecting rabbit semen. J. Reprod. Fert. 7:401-403.

Brun, J.M. et al 2002. The relationship between rabbit semen characteristics and reproductive performance after artificial insemination. Anim. Reprod. Sci 70:139-149

Foote, R.H. and Carney E.W. 2000.The rabbit as a model for reproductive and developmental toxicity studies 

International rabbit reproduction group. 2005.Guideline for the handling of rabbit bucks and semen. World rabbit.Sci. 13:71-91

Lavara, R. et al. 2005. Do parameters of seminal quality correlate with the results of on-farm inseminations in rabbits? Theriogenology 64:1130–1141

Mocéa, E. and Vicenteb, J, 2009. Rabbit sperm cryopreservation: A review. Animal Reprod Sci. 110: 1–24

Morell, J.M. 1995. Artificial insemination in rabbits. British Vet. J. 151: 477-487

Mukherjee, D.P. et al 1951.The Gelatinous Mass in Rabbit Semen  Nature 168: 422 - 423

Theau-Clément, M. 2016. Relationships between rabbit semen characteristics and fertilising ability after insemination. Animal. 10:426-431

Vincente, J.S. et al. 2008. Rabbit reproductive performance after insemination with buserelin acetate extender. Livestock Science 115: 153–157 

Viudes-de-Castro, M.P.Ovulation induced by mucosa vaginal absorption of buserelin and triptorelin in rabbit 68: 1031-1036. Theriogenology 68: 1031–1036

http://placentation.ucsd.edu/rabbitfs.htm Kurt Benirschke.