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Chapter
The Incidence of Ovulation and
Detection of Genes Associated
with Ovulation and Twinning
Rates in Livestock
Ozden Cobanoglu
Abstract
Cattle is a monotocous species that generally produce only one offspring per conception. However, multiple ovulations are a naturally emerging reproduc- tive phenomenon typically controlled by genetic structure and environmental factors. On the other hand, few genes or causative mutations might explain significant genetic variations between animals for the reproductive traits. Studies report different methods, including QTL analysis, fine mapping, GWAS, and MAS selection, to improve such traits due to their economic importance. The recent fine-mapping study, which narrows the genomic region, indeed, influenc- ing multiple ovulation, gives positive signals that causative mutation controlling high ovulation rate may be identified shortly. In conclusion, identifying the major genes that considerably affect ovulation and twinning rates provides the opportunity to increase reproduction efficiency by improving genetic gain in livestock species.
Keywords: ovulation rate, twinning rate, polymorphism, QTL, MAS, livestock
1. Introduction
Complex traits are typically influenced by multiple genes by their combined contributions and modifications of environmental factors. However, a few genes or loci account for most variation between individuals for any given domesti- cated species. Researchers develop various methods, such as marker-assisted selection (MAS), to improve production and reproduction, and performance traits because of their economic significance in dairy and beef cattle over the last 50 years. This chapter presents issues about the major traits with economic values for the genetic improvement of livestock reproduction. It also covers aspects from basic information about physiological mechanisms of ovarian follicular development in ruminants to incidence of multiple ovulations to the fundamen- tal studies of ovulation rate in model species to all aspects of ovulation rates and genetic studies to identify quantitative trait loci or causative mutations affecting ovulation rates and more explicitly twinning rates in bovine species.
The Incidence of Ovulation and Detection of Genes Associated with Ovulation and Twinning... DOI: dx.doi/10.5772/intechopen.
Livestock species are mainly classified either as monotocous species, like cattle, water buffaloes, and horses, or multiparous species, like goat, sheep, and pig based on ovulation rate depending on the characteristic of a species [13]. Biological factors for the consistent multiple ovulations and how to improve or control the ovulation rate in other single-ovulating species in livestock have been of interest to some researchers to understand and intervene in the process of follicular development by applying assisted reproductive technologies. Therefore, identifying various experimental animal models with multiple ovulation rates could efficiently enhance the selection response in farm animals. Specifically, the reproduction process is primarily influenced by genetic and environmental factors for a transformation from primordial follicle to mature ovulatory stage and typically has low to medium inheritance; thus, traditional phenotype-based selection methods are often time-consuming processes due to a lack of efficiency. It is more effective to select breeding animals based on their genotypic structure to increase ovulation rate, prolificacy, and litter size as reproductive abilities in livestock species. Eventually, selecting animals based upon highly polymorphic marker information for reproduction efficiency (MAS) will be of great importance for future breeding programs in the livestock production system.
3. The use of molecular genetic markers and techniques to improve
reproductive performance in livestock
Genetic improvement of reproductive efficiency is one of the most effective strategies available to improve the performance of farm animals. Especially in the last 50 years, selection program based on classical or molecular genetic principles has led to significant positive changes in dairy and beef cattle [ 14 ]. Reproductive efficiency is influenced mainly by environmental factors such as dietary regimen, animal health and management, and their interactions, as well as by many genes in dairy animals. Reproductive traits generally have low-to-moderate heritability and do not show excellent progression to phenotype-dependent selection by classic selection methods. Therefore, determining the genes that affect the reproductive ability and including them in the selection program is one of the crucial arguments in increasing the efficiency and success of the selection process. Genetic markers of follicle number in cattle ovaries can identify heifers that will become highly fertile cows because genes play an active role not only in the physical structure of an organism but also in its functioning. Therefore, analysis of the farm animal genomes will enable us to identify putative genes that are supposed to affect fertility and cow productivity, which are economically important traits in livestock, as the ultimate goal. Specific chromosomal regions, which contribute to complex traits, are called quantitative trait loci (QTL). Several studies were conducted to identify genetic variation in quantitative traits in livestock and laboratory species since the genetic variation is an essential part of breeding programs. A possibility of detecting loci that affect quantitative traits using genetic markers has been realized since Sax’s study with beans, which utilized seed-coat characters as markers due to the relationship with seed size in 1923 [ 15 ]. Selecting desirable alleles at particular loci based on marker information will increase the selection response for the next generation. Short sequences of DNA, called genetic markers, are specific DNA regions in the animal genome that indicate variation within the population. These polymorphic regions can be positively or
Bovine Science
negatively associated with particular reproductive traits of interest. One of the main tools for genetic improvement is the wide usage of molecular markers such as microsatellites, minisatellites, and single nucleotide polymorphisms (SNPs) using different methods such as PCR-RFLP, SSCP, SSR, qRT-PCR, and whole-genome analysis or the next-generation sequencing [ 16 ]. Especially microsatellite markers are not only highly polymorphic but also reason- ably abundant throughout the genome [17]. The relationship between marker alleles and phenotypic observations on the trait is used widely in linkage analysis to map a segregating QTL in a population. The presence of highly polymorphic DNA markers in genetic maps in various farm animals and their relationship to phenotypes provide an effective tool for QTL affecting traits. However, identifying markers closely linked to the target region and determining the association between marker allele and QTL allele, which control the quantitative traits, are rather complex processes. A high-reso- lution marker map and precisely recorded phenotypic values are needed to determine the linkage between marker loci and QTL with low to moderate effect controlling the traits like reproductive performance [ 18 ]. Therefore, the QTL region affecting mainly low-moderate heritable traits is detected to find molecular markers that can be applied in the MAS system, enhancing the genetic gain for the reproductive trait of interest. Several reproductive traits have been associated with fertility in dairy cattle, including age at puberty, early ovulation, size of ovulatory follicles, multiple ovulation, ovarian cystic structure, embryo survival, and heat detection [19, 20 ], which heritability rates are around 1–5% [21]. The prediction of these heritability ratios still notifies that there is a potential to make genetic progress selecting against reproductive traits in bovine. Genome-wide association studies (GWAS) are widely used powerful techniques to discover genetic variants strongly associated with vari- ous complex traits concerning any disease resistance, productive and reproductive abilities in different organisms over the last twenty years. For this purpose, chip- based microarray technology has been developed as a high processing platform to support GWAS analysis. GWAS is a technique that assays high-density SNP markers located throughout the genome to identify putative locations, either causative or in linkage with continuous phenotypic variation. The availability of millions of SNPs markers makes the system easily genotyping on throughput platforms by covering the whole genome [ 22 ]. Various GWAS studies have been carried out on livestock, especially in dairy cattle [ 23 ], beef cattle [ 24 ], water buffalo [ 25 ], pigs [ 26 ], sheep [27], and goat breeds [ 28 ]. However, the large number of potential genes identi- fied by GWAS have not been fully validated yet. As the best-powered studies, they are combining researches of GWAS data and genomic selection (GS) with MAS in livestock species will precisely accelerate the accomplishment of analyzing massive genotypic data through millions of genetic markers which are collected from up to hundreds of thousands of phenotyped animals with diseases and traits of interest soon [29, 30 ]. In addition, other new technologies, including RNA-sequencing technology, to be implemented through the genome-wide sequencing of mRNAs in animal species can be widely applied in such studies over time [31]. In conclusion, it is expected that many more major genes, causative mutations, and even several genes with minor effects will be definitively identified shortly due to the drastic decrease in prices for SNP genotyping and DNA and mRNA sequencing with the substantial increase in livestock genomic studies.
4. Developmental stages of ovarian follicles
Folliculogenesis, the complex biological process of forming ovulatory follicles among the cohort of growing primordial follicles on the ovaries produced by
Bovine Science
as it has been observed in this range in many studies using both Bos taurus and Bos indicus breeds [ 46 –48]. However, the high progesterone concentration prevents the first dominant follicle from maturing, as the corpus luteum has not regressed yet. Thus, the first dominant follicle cannot be functional and ovulatory. Subsequently, a second ovulatory wave can be observed. The dominant follicle from this wave can keep on growing and ovulating during the corpora lutea (CL) regression. In addition, a third source of the ovulatory follicle becomes apparent on day 16 after ovulation in some cattle breeds due to the regression of the second dominant follicle during luteolysis. Even if each wave involves simultaneous emergence of a cohort of follicles, usually one of them, sometimes two, become dominant follicle(s), and all of the others eventually become subordinates. A single oocyte is released from the dominant follicle due to either naturally occurring or artificially induced ovulation. On the other hand, subordinate follicles begin to regress right after a short growing phase [ 44 , 45]. It was noticed from individual to individual that the follicle size at ovulation was quite different. Dairy heifers showing two-wave cycles have a follicle at a diameter of 16 mm in ovulation. However, follicle size is smaller in heifers (13 mm) with three-wave cycles [ 44 ]. Similarly, the size of ovulation follicles has been reported as 14 mm in Holstein heifers. However, the follicle size observed for lactating dairy cows was slightly larger and was found to be 17 mm [ 49 ]. In many studies of fol- licular diameter deviation, both the future dominant follicle and the most signifi- cant lower follicle were more prominent in Bos taurus. However, diameter deviation occurred at similar times after wave emergence in Bos taurus and Bos indicus. Bos indicus has a smaller follicle size when the deviation in the follicle diameter cannot be fully revealed. Nevertheless, the results of the studies support the idea that the future dominant follicle generally has a size advantage over the largest subordinate one [ 46 , 48, 50]. Reproductive biotechnology has recently emerged as a powerful tool to improve livestock productivity and reproductive performance. Therefore, these modern reproductive technologies have started to be used instead of conventional classical techniques in many reproductive-based studies recently. Progress in our under- standing of follicle development and selection has sparked the development of synchronization protocols for fixed-time artificial insemination (AI), in addition to the applications of other cutting-edge reproductive technologies such as in-vitro fertilization (IVF), embryo culturing and transfer (ET), cloning, estrus synchroni- zation, transgenesis, and much more new emerging reproductive biotechnologies [51, 52]. As a result, these developments in terms of sustainable livestock produc- tivity are important for optimal follicle growth and making the right choices to increase reproductive efficiency in livestock species.
5. The incidence of multiple ovulations
Cattle are a uniparous species that means females usually produce only one progeny per conception due to the single dominant follicle in each ovulatory cycle. Alterations in follicle selection can lead to codominant follicles and multiple ovula- tions, which are the basis for multiple births in cattle and sheep [ 53 ]. In rare cases, the synchronous emergence of two follicles as a physiological pattern, albeit in a monovular type, is altered so that the follicle selection mechanism allows both to be selected as the dominant follicle among several follicles in the follicular wave. The ease of evaluating follicular events by trans-rectal ultrasonography and the accu- racy of the data obtained from these studies have allowed cows to be widely used as an ideal research model in follicular studies in ruminants and humans [ 54 , 55].
The Incidence of Ovulation and Detection of Genes Associated with Ovulation and Twinning... DOI: dx.doi/10.5772/intechopen.
Ultimately two oocytes are released from codominant follicles at the end of ovulation due to either natural stimulation or artificial inducements. In the devel- opment of codominant follicles, deviations occur in the diameter of the follicles when the largest follicle and the second-largest follicle are close to 8 mm. The third-largest follicle has a low growth performance, and the deviation in 2nd largest follicle may occur 36–50 h after the deviation of the first follicle [56]. Ovulation of two future dominant follicles occurs either from the same ovary simultaneously, or each follicle consists of a separate ovary [ 57 ]. In a study, synchronous production of two oocytes from different follicles was observed due to the evidence of two corpus luteum (CL) on the ovaries of cattle [ 58 ]. Also, research about follicular develop- ment during the estrous cycle in twin-calving cows indicated that double and triple ovulations coincide from different ovulatory follicles of the same follicular wave rather than ovulation of single mature follicles from two consecutive waves [ 59 ]. In addition, the authors noted that the cysts in the ovary and lack of CL possibly increased the incidence of double ovulation during pregnancy [ 60 ]. Therefore, as more than one follicle deviates and becomes dominant, the chance will be increased for ovulation of more follicles simultaneously. After all, twins, triplets, or overall multiple births in rare circumstances will become a reality if all subsequent events commonly occur for both oocytes from fertilization to parturition. The natural incidence of twin or triplets birth in cattle is mainly due to mul- tiple ovulations that have been summarized in many studies, even if the results are inconsistent [ 61 ]. While the multiple births five decades ago is around 1–5% depending on breed, genetics, parity, and other environmental conditions [62], this rate has increased up to 10–22% in lactating dairy cows today. There have been many studies conducted about regulating multiple birth rates in cattle by select- ing genetically favorable animals [ 63 ], utilizing hormonal treatments [64], or utilizing embryo transfer techniques [ 65 ]. One of the reasons affecting ovulation rates is low progesterone secretion in older cows, which might be the main reason for the increase in circulating LH level and eventually causes enhance ovulation numbers as progesterone has a suppressive effect on LH release. In addition, growth hormone and nutritional treatments greatly influence a multi-ovulation response of an individual [ 66 ]. Also, the ovulations of two follicles simultaneously caused to increase in days of milk among pregnant animals [ 60 ]. In a recent study, the incidence of multiple ovulation rates in early lactating animals was 14%, but they did not significantly affect various reproductive outcomes of cattle [ 67 ]. Although the underlying mechanisms of multiple ovulation have been studied extensively, the dynamics of the entire mechanism are still not fully explained. Monozygotic twins are genetically and physically identical since they are formed from one fertilized egg, splitting into two identical halves during early embryonic developmental stages. Thereby both individuals are always the same sex. In the case of dizygotic or fraternal twins, two different sperm fertilize two completely differ- ent ova simultaneously. Thus the successful result of ovulation and fertilization of two oocytes will be dizygotic twins. Dizygotic twins are not identical genetically or phenotypically as monozygotic twins are. They are not necessarily the same sex as opposed to monozygotic twins. They can also be similar or different from siblings born from the same parents during different gestations [68]. Twin or triplets birth is an unavoidable issue in dairy and beef cattle produc- tion systems which negatively affects the health, production, reproduction and overall decreases the productive life span of animals [ 69 ]. The study reports that the calf survival incidence from twin- and triplet-producing animals were relatively low, around 44% depending on the breed composition [ 70 ]. In a recent study of the economic analysis of multiple births, the economic loss to the livestock breeder from each twin calving was estimated at between $59 and $161 in cow-calf
The Incidence of Ovulation and Detection of Genes Associated with Ovulation and Twinning... DOI: dx.doi/10.5772/intechopen.
populations. If ewes are heterozygotes for any of them, it causes to increase ovula- tion rate to two-three ova. However, if sheep is homozygous for Booroola mutation, it dramatically raises ovulation rate from 5 to 14 [ 83 ]. Another study investigat- ing the ovulation rate records obtained from daughters of ewes inseminated by Coopworth rams to understand the inheritance pattern of ovulation rate also proved that there was another maternally inherited gene affecting productivity traits located on the X chromosome (FecX2w). But the location of this gene is entirely different from the gene on Inverdale FecX locus [ 80 ]. The findings of studies con- ducted about four decades ago have guided many subsequent types of research on this subject, in which sheep are extensively used as model organisms in this subject. Currently, the segregation of five major genes that affect ovulation rate and prolificacy has been characterized at the molecular level in various sheep and goat breeds that cause significant phenotypic variations. Overall the detected genes are bone morphogenetic protein receptor, type 1B (BMPR1B; in Booroola Merino, Javanese, Small Tail Han, Hu, Garole, and Kendrapada breeds) [78, 84 , 85], bone morphogenetic protein 15 (BMP15; in Inverdale, Hanna, Romney, Belclare, Cambridge, Galway, Lacaune, Raza Aragonesa, Olkuska, and Grivette breeds) [82], growth differentiation factor 9 (GDF9; in Belclare, Cambridge, Icelandic Thoka, Santa Ines, Embrapa, Finnish Landrace, Norwegian White Sheep, Ile de France, and Baluchi breeds) [ 86 ], beta-1,4-N-acetyl-galactosaminyl transferase 2 (B4GALNT2 in Lacaune) [ 87 ], and leptin receptor (LEPR in Davisdale sheep) [88]. Causative polymorphism studies in different prolific sheep breeds showed at least 12 identified allelic variants for the BMP15, BMPR1B, and GDF9 genes encompassed in the transforming growth factor-beta (TGF-β) signaling pathway secreted from the oocyte. TGF-β is significantly associated with ovulation rate, lit- ter size, and prolificacy and thus plays a critical role in the folliculogenesis of small ruminants. Many studies reported that the mutations in TGF-β pathway-related genes enhanced ovulation rate (35–100%) in heterozygous animals [89]. Moreover, even if causative mutations for fecundity are not fully discovered, two other genetic variants were identified as FecX2W [ 90 ] and FecD [ 91 ], which are segregated in prolific sheep breeds in recent studies. Similarly, about 20 different candidate genes, including TGF-β related genes, were also detected to play a crucial role in regulating folliculogenesis and prolificacy-related traits in several goat breeds [ 28 ]. To improve the genetic makeup of animals affecting high productivity in livestock, over 30 small ruminants, mostly high-yielding sheep and few goat breeds, have been actively used in candidate gene studies that focused on detecting variation related to reproductive performance-related traits.
7. Ovulation rate studies in bovine
7 Cattle as a model animal for multiple ovulation
As a uniparous species, cattle produce only one progeny in most cases, resulting from ovulation of a single follicle during the pregnancy. Nevertheless, the natural incidence of twin or triplet calving in cattle is mainly due to multiple follicular ovulations concerning breed differences, age of dam, parity, season, the effects of feeding and management systems, geographic location of raised animals, and other environmental effects [62]. Specifically, the incidence of double birth was observed as approximately 1% in beef cattle [ 92 ]. In comparison, this rate was determined as 4–5% on average, ranging from about 1% for heifers to nearly 10% for older cows in dairy cattle [ 93 , 94]. Several studies were conducted concerning underlying causes of multiple ovulation rates, particularly twinning rates in cattle by selecting
Bovine Science
genetically highly polymorphic animals [ 63 ], using trans-rectal ultrasonography or quantifying by circulating AMH concentrations, utilizing embryo transfer tech- niques [ 65 ], or utilizing hormonal treatments [64]. The ovulation rate is closely related to the twinning rate in cattle due to the high genetic correlation between ovulation and twinning rates, ranging from 0 to 1. [95]. Although the genetic control of multiple ovulation in cattle by major genes has long been the subject of research, and there has been significant interest in the mechanism underlying multiple ovulation in bovine species [ 70 ], genes with significant effects on ovulation rate have unfortunately not been identified until recently [96].
7 QTL studies about twinning and ovulation rates
The selection of genetically superior animals in terms of twinning frequency has been practiced in long-term experimental herds in different countries. For this purpose, various research herds for multiple ovulation studies have been imple- mented to be established in various countries for four decades. These herds were begun to set up in the early eighties to select for increased twinning rates in France [97], Australia [ 76 ], New Zealand [98], and Meat Animal Research Center (MARC) of the USDA-ARS in the USA [ 99 ] to develop effective genetic strategies to improve production efficiency including twinning and ovulation rates, meat quality, and animal health in dairy and beef cattle production. MARC twinning population initiated with a total of 307 well-suited cows from twelve different experimental beef, dairy, and dual purposes breeds to study involved in follicular development and recruitments and identify genes affecting primarily twinning rate; later taken into account of ovulation rate in 1981 [ 63 , 100]. These cows were selected based on their high twinning frequencies. The twinning rate can be defined as sequential events due to ovulation, conception, and embryonic survival [ 101 ]. Sires whose dams were founders of the herd and sires whose daughters had high twinning rates were used for breeding the founder cows. In addition, semen collected from sires that mainly originated from Swedish and Norwegian breeds was used in the project. The founder breeds in the herd were mainly Holstein (18%), Swedish Red and White and Norwegian Red (12%), Swedish Friesian (16%), Pinzgaurer (18%), Simmental (15%), Charolais (5%), Angus and Hereford (8%), and other breed crosses (5%) [ 102 ]. The primary objective of the research was to increase the twinning rate in the herd. Therefore, they selected animals based on twinning performance. However, later on, they also evaluated animals’ ovulation rate records for 8 to 10 estrous cycles since ovulation rate is highly genetically correlated with the twinning rate (0) [ 58 ]. Thus, they used an animal model with multi-trait repeated records to predict breeding values for twinning rates in 1990. By applying this methodology, they were able to use information not only from the individual but also from all available relatives for twinning and ovulation rates. The most significant advantage of using ovulation rate records as an estimator of twinning rate is to reduce generation interval and reduce the number of cows retained for several generations. The estimated twinning rate was about 4% in 1984. But this prediction rose linearly to 35% in 1996 [ 100 ]. In the latest report, all the cows with lower estimated breeding values (EBV) were culled from the herd. Thus herd size was reduced from 750 to 250 cows giving birth annually. The twinning rate then was enhanced from 35% to over 50% annually since 1997 [ 103 , 104]. Many studies have been conducted to identify ovulation rate and twinning rate QTL in different cattle populations. Several genomic regions for putative ovulation rate were detected on BTA7 and 23 [ 105 ], on BTA5, 7, and 19 [ 106 ], on BTA5 [ 107 , 108 ], on BTA7, 10, and 19 [ 109 ], on BTA14 [ 101 ] for ovulation rate using the USDA
Bovine Science
major bovine allele contributing to a high fecundity rate in a family of cattle with triplet calving ability throughout the generations in New Zealand. The possible sce- nario for this situation might be that a gene or set of genes should be segregated as a single copy from a dam (Treble) to some descendants through its son (Trio) for single gene inheritance. Moreover, such a unique gene allele is expected to be segregated as dominant or partially dominant in female animals [ 70 ].
Trait Chr (Appx. location as cM)
Population Positional candidate genes (Chr)
Method References
Ovulation rate
7 (40), 23 (27) MARC Twinner CYP21 (23) Interval Map. [105]
Ovulation rate
5 (107), 7(5, 57) 19 (65)
MARC Twinner Interval Map. [106]
Ovulation rate
5 (40) MARC Twinner Interval Map, Assoc./LA
[107, 108]
Ovulation rate
7 (30), 10 (75), 19 (65)
MARC Twinner AMH (7), ESR (10), IGFBP (19)
Interval Map. [109]
Ovulation rate
14 (61) MARC Twinner Interval Map. [101]
Twinning rate
5 (68), 7 (108), 12 (9), 23 (30)
Norwegian Cattle IGF1 (5), CYP21 (23)
Interval Map. [111]
Twinning rate
5 (64) Norwegian Cattle MGF LDLA [112]
Twinning rate
5 (68) US Holstein IGF1 (5) Interval Map./ LDLA
[113, 115 ]
Twinning rate
8 (117), 10 (41), 14 (68)
US Holstein Interval Map. [114]
Twinning rate
6 (55), 7 (25), 23 (67)
Israel Holstein AMH (7), CYP21 (23)
G WA S [116]
Twinning rate
10 (49), 20 (27), 28 (8)
INRA Twin LDLA [117]
Twinning rate
4 (44), 5 (67), 6 (8,44), 7 (68,76), 8 (58), 9 (34), 11 (47), 14 (21, 38), 15 (23), 23 (51), 28 (9)
US Holstein IGF1 (5) LDLA/GWAS [118, 119]
Twinning rate
10 (14) MARC Twinner SMAD3 , SMAD6 , IQCH (10)
Linkage/Fine Map.
[120]
Twinning rate
24 (40) Italian Maremmana
ARHGAP8 , TMEM200C (24)
G WA S [121]
Multiple birth rate
11 (31) Swiss Holstein and Simmental Cattle
LHCGR , FSHR (11)
G WA S [122]
Tabl e 1. Chromosomal locations of quantitative trait loci (QTL) and single nucleotide polymorphisms (SNP) associated with ovulation rate, twinning rate, and multiple birth rate in various cattle breeds [89].
The Incidence of Ovulation and Detection of Genes Associated with Ovulation and Twinning... DOI: dx.doi/10.5772/intechopen.
Several daughters (131) of Trio were born by AI in the USA by following the importation of his sperms at a University of Wisconsin (UW)-Madison research farm from 2008 to 2011. The research reports that a significant bovine allele for high ovulation was identified and mapped on a 2-Mb window on BTA10 (+1. CL per cycle for carriers vs. noncarriers for the marker allele of the high ovula- tion rate) by using fine mapping techniques employed the animals raised at UW-Madison research farm [ 120 ]. Thus, the daughters of Trio proved that there was evidence of a high-fecundity allele transmitting on BTA10 that had a major influence on multiple ovulations in cattle [96]. The detected location was not overlapped with any major genes previously reported for the high ovulation rate and litter size in prolific sheep breeds. Eventually, in addition to the noteworthy reproductive performance of Treble, all of her descendants, including Trio, also displayed extraordinary reproductive performance. Therefore, the members of the Treble family with highly reproductive ability should be heavily employed in gene mapping studies to discover major genes with high fecundity rates. It can provide a significant resource for the subsequent investigation of genetic diversity in bovine productivity [ 70 ]. In the follow-up study, the location of a major gene for high ovulation rate was strongly detected at 1 Mb region of BTA10 using half- sib daughters sired by a bull that assumed to be carriers of the Trio allele due to a single mutation. It is noteworthy that the novel region obtained does not overlap with any major gene previously reported, which significantly affecting ovulation rates in ruminants. Thus, the study reports that the newly identified regions could be employed to track inheritance patterns for multiple ovulation rates using from the carrier father’s lineage since the screening of the aforementioned candidate gene consist of any functionally putative causative mutations in the coding region and 5′ or 3′ flanking regions, reminding that the polymorphic SNP region might affect the expression level of any candidate gene controlling the high reproduc- tive performance of animals [96]. When the follicular and hormonal dynamics of animals carrying the high prolific Trio alleles were examined in animals raised at UW, the Trio carrier animals displayed multiple ovulation. The carriers produced more dominant and ovulating follicles with smaller diameters and volumes in this process due to the slower follicle growth rate close to the beginning of deviation during the entire follicular wave. In the study, even if the deviation times were similar between heterozygous bearing allele from Trio and half-sibling noncarri- ers, a significant increase in the selected number of multiple dominant ovulatory follicles in cow having Trio allele was reported to be associated with the enhanced concentration of FSH secretion close to the deviation time in the follicle. There was also evidence that smaller-sized follicles had more LH receptors in animals carry- ing the Trio allele than noncarriers, supporting the potential novel physiological mechanisms causing the production of multiple ovulatory follicles in the Trio allele carriers [89]. This newly identified candidate region covering 1 Mb in BTA10 contains seven protein-coding genes, of which three of them might be taken into account as puta- tive candidate genes. These genes are the small-mothers against decapentaplegic (SMAD) family member 3 (SMAD3), SMAD family member 6 (SMAD6), those of which are the primary signal transducers for the receptors of the transforming growth factor-β (TGFβ)/Bone Morphogenic Protein (BMP) superfamily ligands [123], and IQ motif containing H (IQCH), which is strongly related with the first menstrual cycle in human females [ 124 ]. The other follow study stated well- conserved SMAD6 gene, which plays a crucial role in preventing the BMP/SMAD- dependent signaling pathway, was 9 times more expressed in carrier animals for the high fecundity Trio allele versus noncarriers using animals in UW-Madison research farm by applying quantitative real-time PCR technique.
The Incidence of Ovulation and Detection of Genes Associated with Ovulation and Twinning... DOI: dx.doi/10.5772/intechopen.
Author details
Ozden Cobanoglu Department of Genetics, Faculty of Veterinary-Medicine, Bursa Uludag University, Bursa, Turkey
*Address all correspondence to: ocobanoglu@uludag.edu
© 2021 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (creativecommons/licenses/ by/3), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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