European Journal of Scientific Research ISSN 1450-216X Vol.34 No.4 (2009), pp.474-484 EuroJournals Publishing, Inc. 2009 http://www.eurojournals.com/ejsr.

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Behavior at Birth and Anatomo-Histological Changes Studies of Uteri and Ovaries in the Post Partum Phase in Rabbits
Boumahdi Zoubida University of Saad Dahlab Blida, Biotechnology Laboratory of Animal Production Blida, Algeria E-mail: zoubidaboumahdi@yahoo.fr Tel: 00213(0)771696826 Belabbas Rafik National School of Veterinary BP 161, Hacene Badi El Harrach Alger, Algeria Theau-Clment Michle INRA. Station dAmlioration Gntique des Animaux BP 52627-31326 Castanet-Tolosan Cedex France Bolet Grard INRA. Station dAmlioration Gntique des Animaux BP 52627-31326 Castanet-Tolosan Cedex France Brown Peter John School of Veterinary Medicine and Science, University of Nottingham LE12 5RD England Kaidi Rachid University of Saad Dahlab Blida, Biotechnology Laboratory of Animal Production Blida, Algeria Abstract The aim of the work was to study the behavior at birth and ovarian and uterine changes in local Algerian rabbits around the post partum phase. Our study has been conducted on female rabbits of a local primiparous population raised at the Breeding Technical Institute (ITELV) at Baba Ali. A total of 20 females were set up and presented to the males. Eight females were pregnant at the 12th day after mating, constituting thus our group of experimentation. These females have been intensively observed during the last two days prior birth in order to determine exactly the time of birth for every rabbit. For 6 rabbit does, the total parturition time was 10-15 minutes. In two rabbit does, parturition in two times was observed 2 and 12 hours interval. One rabbit doe has abandoned its offspring and another one has eaten them. The rabbits were then sacrificed at different intervals of time from 0 h to 48 h post-partum. Their ovaries and uteri were removed, fixed in 10 % of formalin for histological studies. Anatomical and histological modifications of the ovary as well as the uterus have been studied for every rabbit. These observations have shown that during the first 8 hours post-partum the corpora lutea remain prominent on the surface of the ovary. Their number declined at 16 hours and they were replaced by large number of

Behavior at Birth and Anatomo-Histological Changes Studies of Uteri and Ovaries in the Post Partum Phase in Rabbits follicles. This study has allowed us to determine two cases of pseudo pregnant rabbits at 16h and 40h post partum. This state is confirmed by the presence of decidual tissue and giant cells, which could be explained by the fact that being kept together in the same cage after birth, these rabbits could have been overlapped, which has induced the ovulation. These observations also showed that at birth the endometrial epithelium was almost completely degenerated, and its regeneration has started at 8h post-partum. The uterine involution was complete at 48h. Keywords: Rabbits, pregnancy, uterus, ovaries, corpora lutea, follicle.

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Introduction
There is interest in rabbits as a source of protein because of their prolificacy and facility of breeding. However, in Algeria, attempts of intensive rabbit rearing between 1985 and 1988 have failed, because of the lack of knowledge of the detailed requirements of the animals and the absence of suitable commercial food. Since 1990, work within some breeding Institutes and Universities have enhanced the breeding performance of a local Algerian rabbit breed (Berchiche et al., 2000; Belhadi, 2004; Zerrouki et al., 2005). Rabbits are induced ovulators and generally accept mating within hours after birth; the uterus involutes very quickly and loses more than half of its weight within some hours (Lebas, 1994). Rabbits can thus be simultaneously pregnant and lactating (Fortun-Lamothe and Bolet, 1995). In contrast to most other domestic species, during the entire post-partum period the ovary of the rabbit contains pre-ovulatory follicles (Gosalvez, 1986). Characterisation of the local Algerian rabbit breed has indicated that they are of relatively low prolificacy and weight but tolerates hot summer weather well (Zerrouki et al., 2001, 2005). However, until this day very few data are available on the study of the physiology of the reproduction notably the Algerian local rabbit. Recently, Othmani-Mecif and Benazzoug (2005) studied histological changes in the genital tract of pregnant and non-pregnant animals. The purpose of the present work was to study the behavior at birth and ovarian and uterine changes in local Algerian rabbits during the post partum phase.

Material and Methods

Animals Twenty primiparous rabbit does were used for that study. They were between 5 and 6 months old coming from an Algerian local population, which had been bred in captivity over a period of 5 years at the Breeding Technical Institute (ITELV) at Baba Ali, Algiers. The animals were housed individually in cages, fed, watered ad libitum; and received a daily illumination of 16 hours of light. Experimental design The rabbit does were mated with a buck aged 6 months old; eight of them were recognized as pregnant by abdominal palpation fourteen days after mating. Those eight rabbits were transferred to a laboratory so that the terminal stages of pregnancy, and the time of parturition could be closely monitored. After parturition, the rabbits were separated of their offspring and placed in a large mixed cage; mounting behavior was observed while the female rabbits were being kept together in a cage after giving birth. At these different moments after birth, the eight rabbits were sacrificed: 0 hours, 4 hours, 8 hours, 16 hours, 28 hours, 32 hours, 40 hours, and 48 hours, and the reproductive tracts were immediately exteriorized by a ventral midline abdominal incision. The uterine horns were separated,

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weighed and measured. The ovaries were removed from the tract and weighed separately. The numbers of corpora lutea (assumed to represent the number of eggs ovulated) were counted. The ovaries and uterine horns were then fixed in 10% formalin and routinely processed and embedded in paraffin wax for histology; sections are stained by the haematoxylin and eosin (H&E) procedure (Martoja and Martoja, 1967). The presence of different stages of follicular development, and their relative frequency was assessed by counting them in a series of at least 5 sections of each ovary.

Results
Behavior at birth Total parturition time was 10-15 minutes for rabbits n 1, 2, 3, 6, 7 and 8 (Table.1). Rabbit n7 gave birth to eight kits outside the prepared nest and has however abandoned its offspring. Rabbit n4 appeared to be very nervous and frightened; it gave birth to three live kits and then, 12 hours later, to three dead ones; it subsequently ate its three live offspring. Rabbit n5 gave birth on two occasions over a two hour periods. Seven alive young were delivered initially and then one alive, two hours later. Ovarian changes The weights of the rabbits at parturition ranged from 2.7 to 3.9 kg. The weight of ovaries, the number of corpora lutea (CL) and the reproductive performance of the eight rabbits are given in table 1. Examples of the typical histological features or the different ovarian structures are shown in Photos 1 and 2. The weight of individual ovaries ranged from 0.24 to 0.63g; combined ovarian weights ranged from 0.48 to 1.24g.
Photo 1: Primordial follicles (Pf), primary follicles (pf, oocytes surrounded by a single layer of flattened or cuboidal follicle cells respectively) and a secondary follicle (Sf, a central oocyte surrounded by layers of granulosa cells and theca interna and externa layers) are present at the ovarian periphery. Rabbit 2 (4 hours post partum). H&E X40.

Sf pf

Pf

Behavior at Birth and Anatomo-Histological Changes Studies of Uteri and Ovaries in the Post Partum Phase in Rabbits
Photo 2:

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Two early tertiary follicles (tf, oocytes embedded within a cumulus oophorus and a central fluidfilled antrum), primordial (Pf) and primary follicles (pf) are present at the ovarian periphery. Rabbit 6 (32 hours post partum). H&E X10.

Pf tf pf tf
Table 1:
Rabbit number

Ovaries Weight, number of corpora lutea and litter size at birth

Time after birth Ovary weight (g) Total 0.62 0.79 0.68 0.48 0.75 0.79 0.86 1.24 Right 4 1 4 1 2 0 0 0 Corpora lutea Left 4 7 5 2 1 0 0 0 Total born Total 8 8 9 3 3 0 0 0 8 7 9 6 8 5 8 9

Right Left 1 0h 0.32 0.30 2 4h 0.33 0.46 3 8h 0.32 0.36 4 16 h 0.24 0.24 5 28 h 0.40 0.35 6 32 h 0.38 0.41 7 40 h 0.40 0.46 8 48 h 0.63 0.61 (1) Rabbit does supposed to be pseudopregnant.

P+(1) P+

In the rabbit killed immediately after birth large numbers of primary follicles have been observed; there were smaller numbers of secondary follicles and occasional tertiary follicles. At subsequent periods after birth there were smaller numbers of primary follicles (Figure 1). During the first eight hours post partum, generally the total number of CL (Photo 3) reflected the number of born alive. In the rabbit sacrificed at 4 hours after parturition, the discrepancy between numbers of CL and kits born could be explained by the death of one embryo
Photo 3: Corpus luteum (CL, sheets of large, pale granulosa lutein cells). Rabbit 1 (0 hour post partum). H&E X10.

CL

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Boumahdi Zoubida, Belabbas Rafik, Theau-Clment Michle, Bolet Grard, Brown Peter John and Kaidi Rachid
A ruptured mature tertiary follicle (mtf) observed in a supposed pseudo pregnant rabbit doe. Rabbit 7 (40 hours post partum). H&E X10.

Photo 4:

mtf

Figure 1: Follicular populations in the post partum phase (from 0 to 48h).

25 20 Follicle number 15 10 5 0 0 4 8 16 28 32 40 48 0-48 hours post partum Primary follicles Secondary follicles Tertiary follicles

By 16 hours post partum the numbers of corpora lutea has declined and they were absent by 32 hours. Secondary follicles were generally present in larger numbers in the first 16 hours post partum than at later times. Tertiary follicles were present in only small numbers at any time. A ruptured follicle (Photo 4) was observed in the ovary of a rabbit sacrificed at 40 h post partum. Uterine changes The weights and lengths of the uterine horns of rabbits sacrificed at different moments after birth are shown in table 2.
Table 2:
Rabbit number

Weights and lengths of uterine horns.

Time after birth Rabbit weight (kg) Uterine horns Length (cm) Right 20.0 19.0 14.0 19.0 21.0 10.5 10.6 20.0 Left 20.0 29.0 14.0 21.0 14.0 12.0 14.0 15.0 Total 40.0 48.0 28.0 40.0 35.0 22.5 24.6 35.0 Total born 8 7 9 6 8 5 8 9

Uterine horns Weight (g) Right Left Total 1 0h 3.310 21.1 21.1 42.2 2 4h 3.465 20.0 26.1 46.1 3 8h 3.265 20.0 19.3 39.3 4 16 h 2.743 18.0 21.0 39.0 5 28 h 3.750 28.5 21.3 49.8 6 32 h 3.015 11.4 11.7 23.1 7 40 h 3.850 11.6 18.6 30.2 8 48 h 3.975 13.0 9.4 22.4 (1) Rabbit does supposed to be pseudopregnant.

P+(1) P+

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Differences are most apparent after 28 hours. Up to that period total uterine weights ranged from 39.0 to 49.8g; between 32 and 48 hours, total uterine weights were lower (22.4-30.2g). The difference in uterine weights is also apparent when considered in relation to body weight (Table 2). Between 0 and 28 hours the ratio (uterine weight / body weight) was between 12.07 and 16.6; between 32 and 48 hours the ratio varied between 5.64 and 8.63. Changes in uterine horn length follow a similar pattern, ranging from 14 to 29 cm up to 28 hours post partum and between 10.5 and 20 cm between 32 and 48 hours post partum. An asymmetry between the weights and lengths of the left and right uterine horns is evidenced. Reflect in part, areas of uterine hypertrophy and hyperplasia at sites of implantation of different numbers of embryos. Microscopic examination of sections of the uterus revealed a rapid return to normal histological structure. At the time of the birth there was multifocal erosion (Photo 5) of the uterine epithelium; residual cells were small and basaloid. By 4 h post partum, there was an intense mitotic activity within the epithelial cells of the uterine surface (Photo 6). Moreover, at four hours post partum, the endometrial was oedematous and contained scattered neutrophils. Over the following 24 hours the number of neutrophils, and the degree of stromal oedema, has decreased and, by 32 hours the endometrial stroma was normal. While endometrial glands were initially simple tubular, they become more tortuous by 40 hours post partum. Mitotic activity has been observed 8 hours post partum when the luminal epithelium was composed of cuboidal cells, some of them ciliated After 16 hours post partum the surface epithelium had returned to normal, being composed of tall columnar, ciliated cells; nuclei became progressively basal in position (Photo 7). Occasionally, multinucleated giant cells were present, particularly close to capillaries beneath the epithelial mucous membrane in the rabbits killed 16 and 40 hours post partum (Photo 8).
Photo 5: Multifocal erosion of the uterine epithelium (UE). Endometrium (End).Myometrium (My). Rabbit 1 (0 hour post partum) H&E X 10.

My

UE End

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Boumahdi Zoubida, Belabbas Rafik, Theau-Clment Michle, Bolet Grard, Brown Peter John and Kaidi Rachid
Intense mitotic activity within uterine epithelium (UE) and uterine glands (UG). Rabbit 2 (four hours post partum). H&E X 40.

Photo 6:

.UE

UG

Photo7:

Columnar ciliated cells (CC) within uterine epithelium (UE).Endometrium (End). Rabbit (28 hours post partum). H&E X40.

UE CC End
UE UE

Photo 8:

Occasional multinucleated giant cells (mgc) in the superficial endometrium, sub epithelial capillaries (Ca). Rabbit 7 (40 hours post partum). H&E X40.

mgc Ca mgc

In the rabbit sacrificed at 40 h post partum, we notice the presence of important decidual plaques (Photo 9 ) and numerous multinucleated giant cells which invade the surface of the mucous membrane uterine, while this type of cells is totally absent in the other rabbits. Those rabbits (n4 and 7 in table.1) are considered to be pseudo pregnant.

Behavior at Birth and Anatomo-Histological Changes Studies of Uteri and Ovaries in the Post Partum Phase in Rabbits
Photo 9 :

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Decidual plaque tissue (DP) consists of decidual cells associated with uterine epithelium (UE). Myometrium (My). Rabbit 7 (40 hours post partum). H&E X10

UE DP

My

Discussion
The objective of the present work was firstly, to describe the behavior of the rabbit does at the moment of parturition and secondly, to conduct a macroscopic and microscopic study of ovaries and uterine horns during 48 hours following birth. On Twenty primiparous rabbit does utilized to carry out the present study, only eight of them were recognized as pregnant. The lower fertility depends generally on parity order, in fact, Chmitelin et al. (1990) demonstrated that fertility was linked to parity of the female (> 85%: nulliparous, < 70%: primiparous, multiparous: 78, 6 %). The parturient behavior of most of the rabbits was normal, preparing their nest before birth. The parturition in two time points observed in two rabbits, 2 hours and 12 hours interval respectively, has been previously reported by Lebas (1994). The case of rabbits giving birth outside the nest, while abandoning the offspring, has been observed once. This phenomenon as well as the cannibalism has been also reported by Besselievre (1980) and could probably reflect a response to some type of stress such as digestive upset, which makes it feverish and loose the appetite. Under these conditions the female rabbit may kill and eat the youngs (Hafez, 1975; Sandford and Woodgate, 1980). The cannibalism observed in this study, could be explained by the fact that all rabbits were primiparous, which leads to hyperexitable condition associated with a placentophagy (Harkness and Wagner, 1989) and / or by the stress following an environmental change (from the farm to the lab). The ovarian changes described in the present report show that antral follicles are ready to ovulate just after birth. Nevertheless, their number however was low between 0 and 48 h and never greater than two. It is difficult to interpret the results of the description of the follicular populations at various stages post partum because the measures were made on different rabbits which could be explained by the fact that this study has been conducted on different females at each time point. Hulot and Mariana (1982) also observed a variable numbers of tertiary follicles and pre-ovulatory follicle after parturition. Rodriguez et al. (1984) reported that the first wave of follicular maturation postpartum begins at the end of gestation coinciding with a fall in progesterone levels. Falling progesterone levels (Beyer and Mcdonald, 1973; Elsaesser, 1980; Hudson et al., 1990) in association of oestrogens by antral follicles (Challis et al., 1974; Nicosia et al., 1975) explains the sexual behavior of rabbits in the last days of gestation (Beyer and Rivaud, 1969) when female receptivity is high (Moret, 1980). Fortun et al. (1993) showed that most rabbits were susceptible to mating 12 hours after birth and 77 % of these rabbits became pregnant. The kinetics of the follicular growth in the post partum phase could be investigated further by monitoring sequential changes in hormone levels in individual animals. In rabbit does, ovulation is a neuroendocrine reflex, that is inducted by mating or exogenous hormone administration. In the ovary of unmated or un-stimulated rabbits, therefore, functional CL should not be present and, in circulating plasma, progesterone concentration should remain at basal level. The induction of pseudo pregnancy in two rabbits could be explained by mounting behavior

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between females housed together. In fact, these animals had been seen to be mounted while housed with other females post partum. It is assumed that ovulation was induced by that activity but measurement of progesterone levels would be needed to confirm such changes. Indeed, for these rabbit does, some decidual tissue was found. Moreover, the presence of giant cells in the endometrium associated to the decidual tissue suggests a pseudo pregnancy. Since overlapping was observed for one female, we make the hypothesis that the stimulation had lead to an ovulation, probably 10-12 hours later. Since the oocytes were not fertilized (absence of spermatozoa) CL would secrete progesterone until the luteinisation process. On primiparous does inseminated from 0 to 19 days post partum or 2 days after weaning, Theau-Clment et al. (2000) showed that 20% of these females had 2 CL generations and high levels of progesterone associated with low receptivity (22%) and fertility (3%). More recently Boiti et al. (2006) called this phenomenon the high progesterone syndrome. The decidual tissue has already been described in pregnant rabbit does. Yellowish spots are associated with the uterine mucous membrane, including giant cells and surrounded with decidual cells (Hammond, 1917; Larsen, 1962; Larsen and Davies, 1962). The morphological structures of the uterine epithelium and the decidual cells observed under the light microscope from pseudo pregnant rabbit are similar to those observed in the pregnant New Zealand white rabbits by Merad (1988). Uterine involution is defined as the return of the uterus to its normal weight and size after parturition, allowing implantation of further fertilized eggs in preparation of a new gestation (Badinand, 1981; Franck, 1991). Various aspects of uterine involution have been precisely studied in cattle (Gier and Marion, 1968; Kaidi et al., 1991) which are important in determining the reproductive future of the animal. Changes in the epithelial tissues and the interstitial tissue in the rabbits were similar to those described in cows (Kaidi, 1989) and goats (Krajnicakova et al., 2002). These authors mentioned the important role of macrophages in the uterine involution, while in the rabbits described here the neutrophils were predominant. Immunohistochemical staining could be used to investigate more closely the possible role of macrophages in the uterine involution in the rabbits. In the rat, neutrophils and macrophages are involved, during the first four days following birth (Padykula, 1975). The influx of phagocytic cells is important in removing tissue debris as well as helping protect against bacterial infection after parturition (Borrowski, 2006; Krajnicakova et al., 2002). While similarities between uterine involution in rabbits and other species have been observed, it is noteworthy to mention that uterine involution as well as regeneration occurs within 48h post partum. The speed of uterine involution may reflect the type of placentation, since rabbits in which uterine involution is complete within two days have a discoid placentation (Lebas, 1994). In animals with a diffuse placenta (mare and sow) uterine involution is faster (2 to 3 weeks), than animals with a cotyledonary placenta (ewe, goat, and cow) in which it takes 4 to 5 weeks; uterine involution in animals with a zonary placenta (cats, dogs) is extended to 10 to 12 weeks (Slama et al., 1999).

Conclusion
Studies on reproductive physiology of the local Algerian strain of rabbit are rare. The duration, the progress and behavior of rabbits during the birth were described. The present study carried out on the rabbit of Algerian local population is original, as far as the animal behavior and anatomo-histological studies of uteri and ovaries in the post partum phase are concerned. Study of the post partum ovarian changes revealed that tertiary follicles are present at the time of birth, which is consistent with the high sexual receptivity from the first hours following parturition that is well known in this species. However, the kinetics of the follicular growth deserves to be studied more in the post partum phase on the same rabbits in relation with the hormonal profiles and ultrasound studies. At the uterine level, the histological observations brought to light two successive phases; the first one is degenerative from the birth which leads to the destruction of the uterine epithelium, and the second one is regenerative, with an intensification of mitoses from 4 hours. The regeneration of the uterine epithelium is total at 48 h

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post partum. The changes in the rabbit uterus are also characteristic of a female which can be fertilized very soon after parturition.

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