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��ࡱ�>��	,/����)*+�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������[�	���hbjbj����	4lΐΐ"�,�������NN�#�#�#�#�#�����#�#�#8�#��&T�#����&��(�(�(�(`+`+`+j�l�l�l�l�l�l�$9��۫J����#`+`+`+`+`+���#�#�(�(oe�@<@<@<`+b�#�(�#�(j�@<`+j�@<@<�B^0�`�(����0I����#�60r_$V�{�0���_h%��7N%�H�`�`�6%��#���`+`+@<`+`+`+`+`+����@<`+`+`+��`+`+`+`+��������������������������������������������������������������������%�`+`+`+`+`+`+`+`+`+N	W":	Review Article 

Recent development on potential use of intra-vaginal electrical impedance in female farm animal reproduction- a review

Tefera Yilma (PhD)*

*Department of Clinical Studies, Faculty of Veterinary Medicine and Agriculture,
Addis Ababa University, Ethiopia
Tel. +251-911-886605, P.O. Box 31441, Addis Ababa
E-mail: HYPERLINK "mailto:tefera.yilma@aau.edu.et"tefera.yilma@aau.edu.et

Abstract

Electrical impedance is one of the useful techniques that can be applied for monitoring of reproductive activities such as estrus cycle related changes of the genital tract, pregnancy, pending parturition and postpartum resumption of ovarian luteal cyclicity in female farm animals. This paper reviews research findings that have been accomplished during the past few decades in this area of research. Values of electrical impedance that have been measured in the vagina and HYPERLINK "https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/vulva" \o "Learn more about Vulva"vulva during the estrous cycle show significant variation, not only between organs but also different locations of these reproductive organs and among farm animal species. Findings of several studies indicate that changes in the intra-vaginal and vulvar electrical impedance are associated with the HYPERLINK "https://www.sciencedirect.com/topics/veterinary-science-and-veterinary-medicine/ovulation" \o "Learn more about Ovulation"preovulatory HYPERLINK "https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/luteinizing-hormone" \o "Learn more about Luteinizing Hormone"luteinizing hormone (LH)-peak during the estrus period; low values of intra-vaginal electrical impedance (IEI) coincided with the pre-ovulatory LH-peak at the height of standing heat. Similarly, the changes in the measurement values of electrical impedance significantly correlated with plasma levels of HYPERLINK "https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/progesterone" \o "Learn more about Progesterone"progesterone and estradiol-17( during the pre- and estrus period. Similar relationship was also evident between peri-parturient changes in intra-vaginal electrical impedance and plasma concentrations of estradiol-17� and progesterone. Findings of several studies suggest the potential use of intra-vaginal and vulvar electrical impedance to predict the optimal time for artificial insemination during the estrus period and to diagnose early pregnancy. Some research findings also indicate the possibility of using vaginal and vulvar bio-impedance technique to predict imminent calving, and determine postpartum reproductive disorders as well as resumption of ovarian luteal cyclicity.

Keywords: Electrical impedance, Estrous cycle, Parturition, Pregnancy, Vagina
Introduction

One of the major constraints to good reproductive performance in farm animals� reproduction is low estrus detection. Detection of estrus is often difficult mainly due to factors that can be categorized as environmental (extreme temperatures, housing, changing nutritional regimens) and physiological (silent estrus, anestrus, short periods of estrus, onset of estrus during the night hours). The ovarian cycle that consists of follicular phase and luteal tissue development is associated with histological [1] and electrophysiological [2, 3] changes in the mucus membrane of the reproductive tract of female animals. It has been suggested that these estrous cycle related characteristic changes could be the result of alterations in the plasma concentration of ovarian steroid hormones [1], particularly from elevated level of estrogens [3] during the phase of ovarian follicular development and maturation. Histological and biochemical [4] changes of the reproductive tract that are induced by the ovarian endocrine activity can be monitored by measuring of electrical impedance, the ability of a tissue to resist the flow of externally applied an alternating low electrical current [1, 2, 5]. 
Elevated level of estrogen during the follicular phase of the ovarian cycle induces hydration of vaginal and a vulvar tissue, which in turn increases the electrical conductivity of the vaginal mucus and epithelium [6, 7]. Electrical impedance is shown to be one of the useful techniques that can be applied for monitoring of estrus cycle related events that occur in the genital tract of female animals. Values of intra-vaginal electrical impedance show variations during the estrous cycle, and hence, can be used in determining of the stages estrous cycle [8, 9]. Several authors [8, 10, 11] measured markedly declined impedance values at estrus when compared with values of the diestrus period. In support of this, female animals inseminated with low electrical impedance values had highest pregnancy rate than those inseminated when vaginal electrical impedance is high [12-14]. Hence, intra-vaginal electrical impedance values can be applied as a potential tool for estrus detection [15, 1, 10, 16], determining of early pregnancy [12, 10, 13] and prediction of pending parturition [7, 2] in female animals. Nevertheless, there is considerable variation within and among female animals [17, 11] in addition to cyclic changes, partly due to techniques of measurement and genital inflammatory reactions [18, 19].
This review focuses on recent development of the potential use of electrical impedance and its association with ovarian steroid hormones and LH during different events of female farm animal reproduction.

 Changes in vulvar and intra-vaginal electrical impedance during the estrous cycle 

2.1 Female cattle

Several studies have been conducted to evaluate the potential use of intra-vaginal electrical impedance in determining of the optimal time for artificial insemination (AI) during the estrus period in female cattle [15, 12, 10, 20, 13]. The research activities of [13] that have been conducted to verify its potential aid in the improvement of conception rate after AI generally indicated lower mean values of vaginal mucus electrical impedance at the time of insemination, particularly in pregnant cows when compared with the mean values observed in non-pregnant cows. Such studies supported the potential use of electrical impedance measurements for prediction of the optimal time for AI in female cattle due to the fact that higher pregnancy rate has been achieved in cows with low mean electrical impedance values during the estrus period [13, 14].
Previous studies that were conducted by [21] and [1] focused in their respective works on the variation of vaginal impedance values with respect to the placement of probe and stages of the estrous cycle in cattle. The authors found declined impedance values during the follicular phase of the ovarian cycle that coincided with pre-estrus and estrus period when compared with the diestrus phase of the estrous cycle. Moreover, the findings revealed that vaginal impedance values were lowest and least variable when determined in the ventral part of the anterior vagina [21]. Nevertheless, the time at which the lowest values reached varied within and between animals during the estrus period that limits the universal and indiscriminate use of vaginal impedance in determining the height of behavioral estrus. Therefore, some authors highlighted the importance of conducting measurements of impedance frequently and regularly, combined with subsequent analysis of individual impedance curve profile during the different phases of estrous cycle [14]. 
In the work of [1] that conducted to assess the use of electrical conductivity of vaginal and vulvar tissues for detection of estrus in dairy cows, significantly increased conductivity was observed at estrus when compared to a non-estrus period in both vaginal and vulvar tissues of dairy cows. Consequently, the authors confirmed that both electrical frequencies can satisfactorily characterize the changes in tissue conductivity that is associated with estrus period. Monitoring of the genital tissue Bio-impedance (BI) that measured with constantly implanted electrodes in the vulva has been tested to determine its potential application as an aid to reproductive management in female cattle [19]. The findings in similar study revealed that reproductive events that are characterized by changes in fluid content and electrolyte concentration of the genital tissues could be used in the reproductive management of cows [6], through detecting and quantifying the anticipated changes by genital tissue BI monitoring. The fact that tissue BI is quantitatively inversely related with tissue hydrational status, genital BI monitoring can find its potential application in determining of the optimal insemination time [6, 19], considering that intra-vaginal and vulvar tissue BI declines immensely during the pre- and ovulatory stage of the ovarian cycle. 
The use of intra-vaginal electrical impedance (IEI) in detection of estrus, diagnosing of early pregnancy and its relationship with the size of the dominant follicle in dairy cattle has been investigated by [10]. According to the findings of these authors, the mean diameter of the largest follicle (DLF) was highest at estrus during which the vaginal electrical impedance value inversely correlated with DLF. Similar study has been conducted previously by [22] to validate the use vaginal electrical impedance for potential indicator of follicular maturity and appropriateness for timed artificial insemination in estrus and ovulation synchronized beef cows. They concluded, contrary to the findings of [10] that vaginal electrical impedance measurements are not adequately sensitive to differentiate between females with and without large follicle and thus are unable to serve as an aid for prospective decision in determining timed artificial insemination (TAI) after estrus synchronization of heifers and cows. 

2.2 Female pig

The optimum breeding of pigs under farm conditions is related to resumption of cyclical ovarian activity and occurrence of estrus and ovulation post weaning. The return of the ovaries to their cyclical function after pregnancy and lactation is often accompanied by a number of disorders, which is a significant cause of lowered fertility in pig farm herds [27, 28] apart from inaccurate estrus detection. The conventional methods of estrus detection are therefore increasingly insufficient, particularly as number of animals in a herd increases. This requires the use of combined estrus detection methods to accurately determine the optimal time of breeding of estrous sows [16].
Similar to cattle, several authors have been reported on the potential use of intra-vaginal electrical impedance (IEI) in determining of the optimal time for artificial insemination (AI) during the estrus period in female pig [23, 24, 16, 11].  Generally, variations in the values of IEI were evident during the phases of estrous cycle, where high and low values were measured during the diestrus and estrus period, respectively [25, 5].  In the work of [16] IEI values began to decline, right after withdrawal of Regumate� administration in estrus synchronized sows, about two to three days before the onset of estrus, where average low values of impedance have been observed within 36 to 52 hours at times of estrus period. IEI values were also associated with psychosexual behavioral changes and changes in the reproductive tract during estrus, where congestion and edema of the vulva and vagina coincided with time of declining values of impedance, during the onset of pre-estrus phase. In accordance with this, variations in absolute vaginal mucus conductivity readings have been observed between and within sows during the different phases of estrous cycle [26]. 
The research activities of [29] focused on the investigation of the influence of parity on the changes of vaginal impedance in post-weaning sows and reported decreased values of vaginal impedance after weaning. According to these authors, vaginal impedance was higher in sows above 6 parities than in sows from parities 1 to 5, concluding that the parity of sows affects the electrical impedance of vaginal mucus [29]. The association of changes in the vaginal electrical impedance of vaginal mucus with the time of ovulation in post-weaning sows has been studied by [26, 30], whereby decreased values of vaginal electrical impedance between few days and hours before ovulation were observed. Increased measurement values were evident before and right after ovulation. Consequently, lowest impedance values failed to correlate with the time of ovulation, indicating that measurements of vaginal electrical impedance cannot be used to accurately predict ovulation in weaned sows [30].

2.3 Ewes and goats 

Low values of vaginal electrical impedance were observed at estrus in ewes [31] where variations in impedance values were also evident in the same ewe at the same stage in different cycles [32]. The findings of this study also revealed negative correlation between follicular diameter and impedance values, where decreased measurement values were observed in the presence of follicles with large diameter. The authors concluded that impedance measurements should be conducted regularly for individual ewes, combined with subsequent analysis of individual curve profile to detect the estrus period [32].
Investigations were conducted on the changes of vaginal and vestibular electrical impedance during the estrous cycle in ewes [33, 34] and dairy goats [35, 3, 36]. In the work of [34] that performed in ewes, the relationship between electrical impedance values of the cervical mucus and serum estradiol -17� and progesterone concentrations and their possible association with fertility at mating, after estrus synchronization and during the anestrus and estrus period has been evaluated. The research findings of these authors revealed a positive relationship between the electrical impedance values of the cervical mucus and blood serum progesterone concentrations during both estrus and anestrus periods. In support of this, it has been suggested that measurements of electrical impedance of the cervical mucus can potentially be used for the detection of the optimal time for mating after estrus synchronization in ewes [33]. 
In the study that has been carried out in cycling dairy goats, vaginal impedance measured at the anterior vagina, near the cervix showed gradual decrease from pre-estrus to the onset of estrus which then increased after the end of manifestation of estrus [35, 36]. Therefore, the authors concluded that vaginal electrical impedance display changes that are closely associated to estrus related behavioral changes during the estrous cycle in female goats. 

 Mare

Recently, investigation has been conducted on the electrical impedance of the reproductive tracts (vagina and uterine endometrial tissues) and the expression of mucus-related genes to identify the stage of the estrous cycle in mares [37]. The research finding of the authors showed decreased vaginal impedance values during the follicular phase of the estrous cycle in mares, particularly at time of approaching ovulation. It is also suggested that electrical impedance of the uterine endometrial tissues can potentially be used to verify the presence of active corpus luteum (CL) in horse reproduction, due to decreased electrical impedance values in the endometrial tissues during the regression phase of the corpus luteum [37].
2.5. Buffalo cows

The achievement of early cyclicity and pregnancy after calving is of vital importance for sustainable buffalo farms, for which proper detection of estrus and correct time of AI is necessary [38]. In line with this monitoring of vaginal electrical impedance has been conducted during postpartum period in conjunction with the estrus and ovulation synchronization protocol in buffalo cows. Similar to other female farm animal species, decreased vaginal electrical impedance values were evident during estrus, particularly during the second injection of GnRH of the OVSYNCH protocol [38]. Measurement values of vaginal impedance during the follicular phase of the ovarian cycle in the work of these authors suggest its potential use as an indicator of estrus in buffalo cows where daily impedance measurements are recommended in order to confirm the stages of the estrous cycle.
Similar works are available on the use of vaginal electrical impedance for determining of the optimal insemination time during estrus and monitoring of the resumption of ovarian activity postpartum in buffalo cows [39]. In the research findings of these authors, more pronounced decrease in vaginal impedance was evident from diestrus to estrus, where measurement values of impedance started rising after ovulation showing a distinct relationship between vaginal impedance and ovulation.  Hence, the possibility of detecting more estrous buffalo cows was suggested by using vaginal electrical impedance than visual method of estrus detection. In support of this, the finding of the study emphasizes the successful use of vaginal impedance to predict the stage of estrous cycle, ovarian status and ovulation that can potentially contribute to improve conception rates in buffaloes [39].



3. Relationships of intravaginal impedance with serum levels of ovarian steroids and LH during the estrous cycle in female farm animals

Extensive studies have been conducted concerning the relationship between intravaginal electrical impedance and plasma hormonal parameters during estrous cycle in cows, [4, 1, 7], sows [23, 16, 11], ewes [33, 34], camel [40] and buffaloes [39]. It has been reported by [1] that peak concentrations of luteinizing hormone, increased in tissue hydration, and patterns of serum ovarian steroid hormone concentrations have been consistent with the occurrence of estrus during the time of elevated tissue electrical conductivity in female cattle [41]. Similarly, findings of several studies that conducted in sows revealed that intra-vaginal electrical impedance (IEI) decreased few days before the LH-peak and remained low during the estrus period that gradually increased after ovulation. Low values of IEI coincided with the pre-ovulatory LH-peak at the height of standing heat in cattle [1, 41] and sows [11]. Fluctuations in the values of IEI were significantly correlated with changes in plasma LH levels during estrus period in sows, where the average interval between the onset of estrus and pre-ovulatory LH-peak and the time span from LH-peak to ovulation were 16 and 30 hours, respectively [11]. Similarly, [20] reported that measurement of vaginal electrical impedance can be used for detection of LH peak during estrus in sows. The authors highlighted more specifically that monitoring of IEI changes provides a more reliable indication of the pre-ovulatory LH peak than detection of estrus in sows. 
The decrease in the values of intra-vaginal electrical impedance that was evident during the follicular phase have been also coincided with low levels of serum progesterone and increasing concentrations of estrogen [23, 16] in sows. The findings in the work of [16] revealed significant correlation between low IEI values and maximum plasma levels of estradiol-17( at estrus. Similarly, significant association was evident between plasma progesterone levels and IEI values during the phases of estrous cycle.
Based on decreased vaginal electrical impedance values during the estrus period and their relationships with estrus related behavioral changes, patterns of plasma level of ovarian steroids and the pre-ovulatory LH peak; it is suggested that the onset of estrus can better be indicated, and hence, the optimal insemination time can be predicted in post weaning sows [11].
Similar research activities have been undertaken to evaluate the relationship between electrical impedance values of the cervical mucus and serum estradiol-17� and progesterone concentrations during the estrous cycle in ewes [34], where positive relation was evident between electrical impedance values of the cervical mucus and serum progesterone concentrations during both estrus and anestrus periods.  
An interesting study has been conducted by [40] to determine the relationship between the size of the dominant follicle, vaginal electrical impedance, sexual receptivity, and serum concentrations of oestradiol-17� and progesterone during the follicular phase in dromedary camel (Camelus dromedarius). The research findings of the authors revealed no significant differences between the mean impedance values in the animals that had follicles as small as 5 10 mm and as large as 16 20 mm. The authors concluded that electrical impedance of the vaginal mucosa has not been a reliable method for predicting the dominant follicular size during the follicular phase in dromedary camels due to the fact that impedance values failed to correlate with the follicular size at all levels of size categories. Contrary to this, the findings showed positive and inverse relations between oestradiol-17� and serum progesterone concentrations with the overall size of the follicles, respectively [40]. 
Moreover, investigation has been conducted to assess the relationship of vaginal electrical impedance curve profile with plasma progesterone levels in buffaloes. The fall in plasma progesterone profiles was synchronous to decreased values of vaginal electrical impedance, where positive correlation was evident to each other [39].  

Changes in vulvar and intra-vaginal impedance during pregnancy, peri-partum period 

Investigations have been carried out on potential application of intra-vaginal electrical impedance to estimate early pregnancy in heifers and cows [12, 10]. Electrical impedance values of vaginal mucosa that measured three weeks after insemination revealed significantly higher values in pregnant than non-pregnant animals, both in heifers and cows. Similar findings have also been reported in the work of [14].  In both pregnant cows and heifers, electrical impedance of vaginal mucosa was significantly higher than non-pregnant animals, and higher than values determined on the day of insemination as well as during the early few days post-insemination. It was also evident that the vaginal impedance measurements were significantly correlated with plasma progesterone concentrations both in pregnant and non-pregnant groups of animals, suggesting that vaginal impedance values can potentially be used as an indicator for the diagnosis of early pregnancy in cows and heifers. Apart from estrus period, low IEI values were also observed at times of approaching parturition in pregnant sows and cattle in the works of [42] and [2], respectively. It is suggested that declining in IEI values during parturition in the works of these authors may be induced by increased plasma level of estrogen similar to estrus period.
Vaginal electrical impedance changes around the time of calving have been studied and described using bipolar electrodes implanted inside the vulvar tissue of pregnant cows [13], where the relationship between peri-parturient changes in impedance and plasma concentrations of  estradiol-17� (E2) and progesterone (P4) has been determined. The research findings of these authors revealed decreased impedance values during few days before parturition and during the initiation of parturition, suggesting that the decrease in impedance values was caused mainly by an increase in the extracellular compartment of the tissues of the genital tract. Decreased concentration of plasma P4 and increased of E2 were evident during the periods of approaching parturition [13]. In support of this, the authors highlighted that impedance values remained low until parturition, while P4 and E2 continued to decrease and increase at calving, respectively.
Increased values of impedance were also evident during the postpartum period that remained high thereafter. To this end, it is suggested that measuring intra-vaginal electrical impedance could be a useful tool for determining of approaching calving; monitoring the recovery of genital tissue after calving as well as studying the dynamics of hormone action on the hydration of tissues of the reproductive tract.  According to [19], vulvar bio-impedance (BI) that measured with constantly implanted electrodes, declines few days before, and remains low until calving indicating that genital BI measurements can be used to detect imminent calving. Conversely, vulvar BI increases gradually during the whole postpartum period and stabilizes at about the time of resumption of ovarian luteal cyclicity [19]. The research findings of these authors also indicated that embryo yield and quality and pregnancy rate is inversely related with the peri-estrous decline of intra-vaginal BI in donor and recipient cows, respectively.

Conclusion

The studies suggest that measurement of intra-vaginal impedance can be used as an indicator of the ovarian status, stages of the estrous cycle and to predict the optimal insemination time during the estrus period as well as to diagnose early pregnancy. Moreover, research findings revealed temporal association among the onset of estrus, estrus related changes in the intra-vaginal impedance values, plasma concentration of ovarian steroids and pre-ovulatory LH peak and ovulation. Some research findings also indicate the possibility of using vaginal and vulvar bio-impedance technique to predict imminent calving, and determine postpartum reproductive disorders as well as resumption of ovarian luteal cyclicity.

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��������������������������������������������������}�i��'hhh�X�0J!5�6�B*CJaJph�-�jhhh�;�0J 5�6�CJUaJhhh�X�0J 5�6�CJaJ'jhhh�X�0J 5�6�CJUaJh�l�h�X�0J 5�CJaJh�MN5�CJaJ$h�l�h�X�0J!5�B*CJaJph� jh�l�h�X�5�CJUaJh�l�h�X�5�CJaJ)����������������PPPP#P/P2P9PHP�P�P�P�P�PQ����ǹǷ�ǘ�}�}�}m}\N= h�l�h�dCJOJPJQJaJh�MNCJOJPJQJaJ h�l�h�X�CJOJPJQJaJh�l�h�X�6�CJOJQJaJh�l�h�X�CJOJQJaJh�MNCJOJQJaJh�l�h�X�5�CJaJ$h�l�h�X�0J!5�B*CJaJph�Uh�l�h�X�0JR5�CJaJ$jh�l�h�X�0JR5�CJUaJhh0J 5�CJaJh�l�h�X�0J 5�CJaJhh0J 5�6�CJaJ10.2478/jvetres-2018-0080
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42. Schams D, Schallenberger E, Hoffmann B, Karg H (1977)Q�Q�Q�Q�Q�Q�Q�Q�Q�R�R�R�R�RSS��Ͼ���{�hX�G9( h�l�h�dCJOJPJQJaJh-CJOJPJQJaJ h�l�h�X�CJOJPJQJaJhh6�B*OJQJ^Jph�$hhh�d6�B*OJQJ^Jph�%h�l�h�d0JHB*OJQJ^Jph�!h�l�h�dB*OJQJ^Jph�h�MNB*OJQJ^Jph�!h�l�h�X�B*OJQJ^Jph� h�l�h�dCJOJPJQJaJh�MNCJOJPJQJaJ#hhh�MN6�CJOJPJQJaJh�l�h�dCJOJQJ\�aJ�P�Q�R	T�T�U�V�XY�Y�h�h�h�h�h��������}�{{{{$
���h���dh�^�h`���a$gd�$�hdh�@&^�ha$gd�$�h���dh�@&^�h`���a$gd�$�h���dh�^�h`���a$gd�$�h���dh^�h`���a$gd�$�h���dh�7$8$H$^�h`���a$gd�SSSSS�S�S	TTTTST[T^TcTdT�T�T�T�T����Ͻ᭝����p^�L?�hhCJOJQJ\�aJ"hhh�d6�CJOJQJ\�aJ"h�l�h�d6�CJOJQJ]�aJh�l�h�dCJOJQJ]�aJh�l�h�dCJOJQJ\�aJh-CJOJQJ\�aJh�l�h�M�CJOJQJ\�aJh�l�h�X�CJOJQJ\�aJ#hhh�d6�CJOJPJQJaJ#h�l�h�dCJOJPJQJ]�aJ h�l�h�dCJOJPJQJaJh-CJOJPJQJaJ�T�T�T�T�TSU�U�U�U�U�U�U�U�U�V�V�V�V���Ͻ����zm�[�I�9hKD�h�dCJOJQJ\�aJ"h�l�h�d6�CJOJQJ]�aJ"h�l�h�d5�CJOJQJ\�aJhKD�CJOJQJ\�aJh-CJOJQJ\�aJh�l�h�X�CJOJQJ\�aJ&hhh�d6�CJOJPJQJ\�aJ#h�l�h�dCJOJPJQJ\�aJ#h�l�h�dCJOJPJQJ]�aJ#h�l�h�X�CJOJPJQJ]�aJh�l�h�dCJOJQJ\�aJh�l�h�dCJOJQJaJ�V�V�V�V�V�V�V�V�V�V�V�V�V�VWWWW#W$WCWDWTWUWWWXWwWxW�W�W�WfXyX|X�X�X�X�X������òò���ò����ò����ò�垇�q`��!hhCJKH$OJPJQJ\�aJ*hhh�d6�CJKH$OJPJQJ\�aJ-h�l�h�d6�CJKH$OJPJQJ\�]�aJ'h�l�h�dCJKH$OJPJQJ\�aJ h�l�h�dCJOJPJQJaJ%jh�l�h�dCJOJQJUaJh�l�h�X�CJOJQJaJh�l�h�dCJOJQJaJhhCJOJQJaJ%�X�X�X�XYYYYYY Y$Y%YjYvYxYyY�Y�Y�Y�Y�Y�Y�Y�����ݹ��������n���_S�Dh-h-CJOJQJaJh-CJOJQJaJh�l�h�X�CJOJQJaJ'h�l�h�d5�CJOJQJaJmHsH'h�l�h�d6�CJOJQJaJmHsH$h�l�h�dCJOJQJaJmHsHh-CJOJQJaJmHsH$h�l�h�X�CJOJQJaJmHsH h�l�h�dCJOJPJQJaJh�l�h�dCJOJQJaJ%jh�l�h�dCJOJQJUaJ�Y�Y�Y�Y�Y�Y�Y�Y�Y�Y�Y�Y�Yh�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h�h����������⻹⭝�⑉�������}y}p}yl��h7$eh�;�mHnHuhY?jhY?Uh�sHjh�sHUh�dCJOJQJaJhhhKD�6�CJOJQJaJhhCJOJQJaJUhKD�CJOJQJaJh-h-CJOJQJaJh-CJOJQJaJh�l�h�dCJOJQJaJh�l�h-CJOJQJaJ$ The oestrous cycle of the cow: Hormonal parameters and time relationships concerning oestrus, ovulation, and electrical resistance of the vaginal mucus. Acta Endocrinologica, 86, 180-192.














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