PJZ_58_4_1911-1925

INTRODUCTION

Enhancing food security through livestock husbandry is of highest priority worldwide. Lately, there is an increasing demand for goose meat for human consumption (Ni et al., 2022). Goose meat is highly preferred as it is rich in niacin, fat, vitamin A, protein, sugar, and vitamin B and considered healthy since it has less cholesterol and low fat but precious in protein (Ibtisham et al., 2017). In garments industry, goose feathers are important inputs to final product for clothing and bedding (Liu et al., 2020). The most reared domestic goose breeds are the well-known species of Anser anser and Anser cygnoides descending from ancestral family Anatidae. Anser anser domesticus is largely raised in Western Asia, Northern Africa, and Europe instigated from the Greylag goose (Kozák, 2019). Anser cygnoides domesticus is mainly bred in eastern Asia, known as indigenous Chinese goose evolving from the Swan goose (Gyuranecz et al., 2020).

Statistically, China is the leading country in goose farming as it produces roughly 95.2% of goose meat, thus 2.4 million tons of 2.5 million tons worldwide annual meat production (Yu et al., 2020). Based on universal percentage proportionality, goose industrialization is still tracking from behind and this is linked to poor egg laying performance. The poor egg laying performance of geese is affiliated with environmental adaptability, nutrition, diseases outbreak, strong broodiness traits, and chiefly prolonged follicle recruitment cycles (Sun et al., 2019; Wang et al., 2021).

Folliculogenesis commences with one follicle being selected from a resting pool of primordial follicles entering prehierarchical, hierarchical, and ovulatory phase while other competitive follicles undergo cell degeneration (Sello et al., 2020). Avian follicle developmental stages are classified according to size, color, and the thickness of follicular membranes (theca and granulosa cells) (Lei et al., 2020). Development and selection of dominant follicle to attain maturity and its readiness to ovulate rely on endocrine signals from pituitary gland and paracrine signals from ovarian follicles and other growth factors (Songsasen and Nagashima, 2020). These orchestrated mechanisms of follicular development are also not functioning in isolation from molecular driven signaling cascades. The nonhierarchical stage of ovarian follicles is the critical phase of the successive fundamental processes to choose one healthy dominant follicle for ovulation per cycle (Liu et al., 2015). Therefore, this stage of folliculogenesis requires extra effort to unravel the growth factors involved, gene profiling, and unique signaling pathways. Currently, the next generation sequencing (NGS) technique is leading in various fields of studies including those interested in goose egg production, offering useful techniques in analyzing gene expression patterns at each stage ovary development (Ouyang et al., 2020; Qin et al., 2021). To our knowledge, there are no studies conducted to summarize the discoveries on the contributary effects and role associated with biological growth factors (Anti müllerian hormone (AMH), bone morphogenic protein 4 (BMP4), small mothers against decapentaplegic homolog 9 (SMAD9), inhibins, melatonin hormone, and heat shock protein 27 (HSP27) during goose ovarian follicular development. This literature review will focus on developmental stages of goose ovarian follicles along with somatic follicular membranes and the phenotypic differences between the two goose species as well as genes and transforming growth factor β (TGF-β) family members associated with follicular developmental stages.

The information used in this review was obtained from scientific research articles published in three databases PubMed, Google Scholar, and ScienceDirect. The general keywords used in search for relevant articles included “goose ovarian follicles”, “gene expression”, “egg production”, “poultry”, “prehierarchical follicles”, “hierarchical follicles”, “AMH”, “BMP4”, “SMAD9”, “inhibin”, “melatonin hormone”, and “HSP27”. The findings of each study were compared to the citations in the discussion sections for comprehensive inclusion of such information to the current review article. These data used in this review were extracted from English written research articles and review articles, however, review articles were mainly used to trace back to the original research articles. The findings about the candidate gene from the kingdom animalia were considered to support the relevance of particular gene in reproduction in comparison to goose. The data provided in this review was not limited to years of publication. In total, this review is comprised of 139 references that provide a comprehensive analysis of the selected topic.

EGG YIELD OF DIFFERENT BREEDS

A number of factors affect goose egg yield such as breed, environment, and age. Table I summarizes the annual performance for individual laying breed. The annual egg production per bird ranges from 25 to 100 eggs.

Table I. Estimated egg yield per annum for different breeds of goose.

Breeds	Species	Distribution by provinces	Annual egg production per bird	Reference
Zi goose	A. cygnoides	Jilin and Heilongjiang Province	80-100 eggs	(Ji et al., 2020; Zhang et al., 2013)
Zhedong goose	A. cygnoides	Zhejiang province	30–40 eggs	(Zhao et al., 2013)
Huoyan goose	A. cygnoides	Liaoning province	70 to 100 eggs	(Wang et al., 2011)
Mangang goose	A. cygnoides	Guangdong Province	30 to 50 eggs	(Qin et al., 2013b)
Sichuan white goose	A. cygnoides	Sichuan province	80 to 100 eggs	(Kang et al., 2014)
Xupu goose	A. cygnoides	Hunan Province	30 eggs	(Qin et al., 2021)
Taihu goose	A. cygnoides	Taihu Lake drainage in eastern China	75 eggs	(Liu et al., 2021)
Shitou goose	A. cygnoides	Raoping County, Guangdong Province	25 eggs	(Liu et al., 2021)
Wanxi white goose	A. cygnoides	Anhui province	25 eggs	(Chen et al., 2012b)
Wanjiang white goose	A. anser	Anhui province	More than 90 eggs	(Chen et al., 2011)

DOMESTIC GOOSE SPECIES

Swan goose (Anser cygnoides)

Anser cygnoides goose species evolved from wild swan goose (Fig. 1A) mainly in Central Asia, Mongolia, South Eastern and Northern China, Southern Russia and wintering in North/South Korea and Eastern and Central China (Wang et al., 2016). Loss of habitat due to unwarranted hunting of swan goose caused a falloff in their distribution and eventually is regarded as endangered large goose species (Randler, 2003). Wild swan goose is recognized by its eloquent plumage with grey stripes, long black bill, and orange legs whereas the domesticated swan goose species all have white plumage with orange legs and bill (Ren et al., 2021). Mostly, A. cygnoides domesticus farmed in Africa (African goose) and China (Chinese goose) is distinguishable by a pronounced protrusion on the frontal area of the skull although obscure in A. cygnoides wild species (Mead, 2013). This protrusion is commonly referred to as a “knob”. The knob size varies relatively within and among the breeds of A. cygnoides congruent to age and sex (Deng et al., 2021). The knob is slightly larger in male than in females and bigger in grownups than adolescents (Ji et al., 2021).

Greylag goose (Anser anser)

The greylag goose (Fig. 1B) belongs to phylum Chordata in the class Aves (Basyouny et al., 2014). Anser anser domesticus goose descended from an ancestor wild goose called greylag (Honka et al., 2018). A. anser is widely distributed in European countries mainly in Ukraine, Hungary, and Poland (Cilavdaroğlu et al., 2020). A. anser goose is characterized by varying color of plumage distributed from cranial to the caudal regions and mostly appears to be brown, grey white or cream/buff with pink or orange legs (Yang et al., 2022). A. anser have either chunky, duck-like or orange long beak and thick neck. A. anser goose on average weighs 2.9-3.7 kg contingent on sex, season, and age, body length ranges between 76-89 cm and wingspan stretching between 147-180 cm (Fox and Kahlert, 2005; Mirzaeinia et al., 2020). This species is large, heavy and show limited flying capabilities (Islam et al., 2016).

BROODING IN GOOSE

Brooding is the caring behaviour displayed by domestic female fowls resulting in temporary termination of oviposition (Jiang et al., 2010). Brooding fowls are overprotective by obstinately sitting on the nest to incubate clutch of eggs, leave the nest for short stretch of time, reduced food and water intake, and the elevated body temperature (Romanov et al., 2002). The cessation of laying is orchestrated by fallopian and ovarian atrophy (He et al., 2022). At this period, the white pre-hierarchical follicles (WPF) are prominent while the yellow pre-hierarchical follicles (YPF) diminish limiting the successive transition of follicle growth/development to ovulation (Liu et al., 2018). The brooding comportment of goose is identical to that of chicken; however, goose has strong brooding characteristics based on the clutch size and total number of eggs laid per mature female per annum (Chen et al., 2014). In birds, folliculogenesis is regulated by number of factors including the external stimulus (light) actuating the endocrine activity triggered by hypothalamic-pituitary-gonadal (HPG) axis (Wang et al., 2019). For instance, light stimulates the hypothalamus to secrete gonadotrophin-releasing hormones (GnRH) to the pituitary gland which synthesize and release follicle stimulating hormone (FSH), prolactin hormone (PRL) and luteinizing hormone (LH) (Fattah et al., 2021). The FSH, PRL, and LH affect the gonads to functionally initiate sex steroid hormone release “estrogens, gestagens, and androgens” and gametogenesis “oogenesis and spermatogenesis” (Luan et al., 2014). The FSH, PRL, and LH have been shown to significantly contribute to folliculogenesis and therefore suggesting their potential role during folliculogenesis.

FUNDAMENTAL UNITS OF OVARIAN FOLLICLE AND SUBSTANTIAL DEVELOPMENT REGULATORS IN GOOSE

Follicular development and ovulation are facilitated by biological crosstalk between the granulosa cells, theca cells, and oocyte through autocrine, endocrine, and paracrine signaling controlled by HPG axis (Li et al., 2021). The anatomy and physiology of mammalian ovaries have been comprehensively studied. However, in the geese, the knowledge about the inter and extracellular factors modulating the follicle viability is inadequate. The corresponding signaling pathways during follicle development trigger heat shock proteins (HSPs), small mothers against decapentaplegic (SMAD) related genes, members of transforming growth factor family inclusive of AMH, bone morphogenic proteins (BMPs), inhibins, as well as the glands secretion products such as melatonin in a stage and time dependent manner reaffirming their indispensable roles in follicle maturation.

Oocyte

The release of a mature healthy oocyte for fertilization is the center for female reproduction process. Oocyte maturation is initiated as soon as the follicle enters the pool of growing follicles and is almost completed upon the antral follicle (Sen and Caiazza, 2013). Before fertilization, multiple intercellular communications controlled by various signaling pathways at different stages of development are essential for gaining the developmental competence of the oocyte. These cellular intercommunications are of crucial importance to maintain the oocyte arrested in the diplotene stage of prophase 1 of meiosis during prenatal life or immediately prior to birth (Fabritius et al., 2011). During follicle development, the ovulatory gonadotropins known as FSH and LH surge stimulate the resumption of meiosis which progress by orchestrated disassembly germinal vesicle (nuclear envelope) termed germinal vesicle breakdown (GVBD) and eventually the oocyte enters the metaphase II (MII) stage until fertilization (Zeleznik, 2004).

Granulosa cells

Follicular growth and development begin with the formation of several layers from the somatic cells, oocyte enlargement and a fluid filled antrum. During the early antral follicle recruitment stage, follicular growth relies dependently on the pituitary gonadotrophins such as FSH (Tiwari et al., 2015). At this stage, the antrum divides the granulosa cells into two separate cell layer parts: the outer layer (mural granulosa cells) and inner layer (cumulus cells) which encircle the oocyte. The mural granulosa cells (MGCs) possess endocrine function and support follicle growth whereas the cumulus granulosa cells (CGC) have a high rate of proliferation, low steroidogenic capacity, low LH receptor (LHR) expression and high levels of insulin growth factor I (IGF-1) (Mehlmann, 2005). Decreased granulosa cell-oocyte crosstalk impedes the transmission of nitric oxide (NO), guanosine 3ʹ,5ʹ-cyclic monophosphate (cGMP), adenosine 3ʹ,5ʹ-cyclic monophosphate (cAMP) levels to the follicular oocyte (Jahromi et al., 2015). Granulosa cells provide nutrients for oocyte maturation through cell-to-cell gap junction intercommunications (Chaube et al., 2005) and their intactness protects oocytes from oxidative stress damage by reactive oxygen species (ROS) (Tiwari et al., 2017). Granulosa cells generally produce sex steroids, and numerous growth factors that interact with the oocyte during its development. The sex steroid production consists of FSH that stimulates granulosa cells to convert androgens produced by thecal cells to estradiol by aromatase during the follicular phase of the reproduction cycle (Magoffin, 2005).

Theca cells

The theca cells are divided into two layers, the theca interna and the theca externa. The theca interna produces androstenedione, and indirectly produces 17β estradiol (E2), thus providing the neighboring granulosa cells with androstenedione, which can be converted into estradiol by enzyme aromatase (Wu et al., 2015). Estradiol has multiple functions on granulosa cells including cell apoptosis inhibition, increasing gonadotrophin expression, promote folliculogenesis, and promotes LH receptors formation on the granulosa cells, which also possesses FSH receptors (Manna et al., 2009). Additionally, the theca interna possesses luteinizing hormone receptors and greatly vascular with no FSH. Theca cells are also a source of keratinocyte growth factor (KGF), a factor that has been shown to suppress apoptosis in cultured preantral follicles. Moreover, theca cells are much differentiated with structural characteristic features of steroid-secreting cells including agranular endoplasmic reticulum containing enzymes necessary to produce androgen; lipid vesicles which store the cholesterol esters (precursors for steroid hormone synthesis) which are transported into the mitochondria by steroidogenic acute regulatory protein; and abundant mitochondria with vesicular cristae which contain the first enzyme in the steroidogenic pathway (cholesterol sidechain cleavage cytochrome P450 (CYP11A)) (Ginther et al., 2003).

Anti-mullerian hormone (AMH)

Anti-Mullerian hormone (AHM), formerly known as Mullerian-inhibiting substance (MIS), belongs to the transformation growth factor beta (TGF-b) superfamily produced by the sertoli cells in male’s testis and granulosa cells in females during ovarian follicular development and growth (Zhao et al., 2018). It is however undisputable that more investigations about AMH focused on mammalian ovary development as it possessed paramount regulatory function during the transition of primordial follicles from the resting pool of follicles to mature ovary ready for ovulation (Nilsson et al., 2011). AMH have been shown to restrict the pre-hierarchical follicles maturation in laying hens (Huang et al., 2021b), inhibit initiation primordial follicles in mouse (Durlinger et al., 2002), bovine (Yang et al., 2017), and human (Carlsson et al., 2006) by deactivating the granulosa cell sensitivity to FSH (Carlsson et al., 2006). In goose, the abundance of AMH was inversely proportional to the thickness of the ovarian granulosa cells as the follicles progress towards maturity (Yang et al., 2019), and less detected in laying goose than brooding goose in time specific manner (Wang et al., 2021). The AMH mRNA levels increased significantly in a stage dependent manner during follicles development and gradually drop to optimum levels over time (Hu et al., 2021). Immunized goose against AMH showed improvement in follicles development rate with an increase in number of eggs per cycle and established the short brooding period (Chen et al., 2020a; Zhang et al., 2021a).

Heat stress and heat shock protein 27 (HSP27)

Internal (physiological) and external (environmental) stress weaken the follicular walls, thereby hindering normal ovarian activities related to growth and development (Abdelnour et al., 2020). Heat stress has lethal effects on the female reproduction performance of animals as it causes the gonadal hormones imbalances and consequently affects oocyte development and quality, thus reducing fertility (Bei et al., 2020; Tu et al., 2016). HSP27 is a member of the p38 MAPK signaling pathway classified as class II type of small heat shock proteins (15-40 kDa) widely distributed in various tissues induced by various intra or extracellular stress stimuli including heat shock, ROS, and death receptor ligation (Bei et al., 2020). HSP27 proteins are involved in multiple cellular functions such as signal transduction and development, cytoskeletal integrity maintenance, protein degradation and folding, cell cycle, cell death differentiation, and stress tolerance (Santana et al., 2020). Previous studies indicated that HSP27 acts an anti-apoptotic chaperon that inhibits the activation of cell death ligands (caspase 9) by obstructing the mitochondrial cytochrome C collaboration with apoptotic protease activating factor (APAF) from arbitrating programmed cell death signaling cascades (Shan et al., 2021). Follicular atresia is referred to as automated follicular cells degeneration mainly the granulosa and theca cells exposing the oocytes to detrimental apoptotic effects initiated by TNF-and Fas death receptor proteins (Sugimoto et al., 2010). HSP27 transcripts or proteins were detectable in normal human oocyte and the polycystic ovarian syndrome (PCOS) and can be associated with oocyte maturation and fitness to reach ovulatory stage (Cai et al., 2013). The mRNA expression patterns of HSP27 in bovine granulosa and theca cells varied with the condition of the follicles and maturity stage (Velazquez et al., 2010). In mice ovaries, the distribution of HSP27 transcripts showed a downward regulation expression over time with the highest level at initial stage of heat shock treatment indicating that HSP27 genes play a protective role from heat stress induced damage (Bei et al., 2020). Although HSP27 has been identified as anti-apoptotic protein, its downregulation in mouse enhanced the oocyte development and maturation and yet improved oocyte apoptosis at early stage by activating extrinsic, caspase 8-aided cascade (Liu et al., 2010). We have for the first time revealed that goose pre-hierarchical follicles convey HSP27 gene during normal ovarian growth/development with predominance in both theca cells and granulosa cells of the middle white follicle suggesting its potential contributary effects during folliculogenesis (Sello et al., 2020). The functional role of HSP27 during the morphological changes related to follicular development/growth and degeneration needs to be explored to understand molecular mechanisms underlying goose folliculogenesis.

Bone morphogenetic protein 4 (BMP4)

Bone morphogenetic proteins (BMPs) are the greatest growth factors, members of TGF-b superfamily functioning in cell differentiation, proliferation, growth, and degeneration (Halloran et al., 2020; Shimasaki et al., 2004). These groups of proteins were first discovered during bone formation (Urist, 1965). The inquisitiveness on BMPs existence further discovered their involvement in autocrine/ paracrine regulation of mammalian ovary development and their abundance vary with cell types and developmental stages (Shimasaki et al., 1999; Shimizu et al., 2004). According to Lankford and Weber (2010), downwards expression of BMP4 was inversely proportional to the stage of ovarian developmental stage in rainbow trout. Furthermore, BMP4 higher abundance was detected in ovarian somatic cell layers than oocytes in Zebrafish ovary (Li and Ge, 2011). Likewise, the BMP4 expression levels varied in chicken granulosa and theca cells preovulatory follicles with exceptional abundance in the granulosa cells of largest follicles (F1) suggesting its possible functional contribution in proximity to ovulation (Onagbesan et al., 2003). BMP4 was reported to enhance prehierarchical follicles resistance to apoptosis and promoted follicles development in chicken granulosa cells exposed to physiological endoplasmic reticulum stress (Yao et al., 2020). Besides the ovary cell types, BMP4 immunoreactions were detected in other reproductive tract organs such as oviduct and uterus throughout estrus stages/cycle in mice (Tanwar and McFarlane, 2011). Contrary to chicken, in goose, fewer amounts of BMP4 transcripts were detected in F1, but predominant in prehierarchical follicles and repressed granulosa cells caspase-mediated cell death through PI3K/AKT/Caspases signaling pathway (Yuan et al., 2019). In some studies, BMP4 inhibited the secretion of progesterone by granulosa cells (Pierre et al., 2004; Spicer et al., 2021) and FSH and FSHb mRNA expression from pituitary cells in ovine (Faure et al., 2005). Detection of BMP4 and its receptor molecules in female reproduction organs provides evidence about their contributive effects in maintaining follicular viability.

Small mothers against decapentaplegic homolog 9 (SMAD9)

The development/growth follicles rely mainly on complex interaction of cell-signaling pathways producing either positive or negative feedback information (de Figueiredo et al., 2018). The SMAD polypeptides are important signal transducers of TGF-b comprising of different types of BMPs, inhibins, activins, and nodal (Coda et al., 2022). The SMAD signaling pathway interaction with BMPs signaling cascades regulate follicular development through binding to distinct forms of receptors (Atwood and Meethal, 2016). According to Chen et al. (2012a) receptor-regulated Smads (R- Smads) downstream activities are phosphorylated by BMP type 1 receptor kinases through ligand binding. In mammals, BMP/Smads signaling cascade played functional roles during the dominant follicle selection, follicular atresia, and development (Costello et al., 2009; Haugen and Johnson, 2010). Smad9 is a member of R- Smads that acts synergistically or antagonistically with BMP signaling pathways depending on its expression levels (Tsukamoto et al., 2014). Interestingly, Smad9 can also be associated with male prolificacy traits as it has been found in reproduction tissues in rams (Chen et al., 2020b). According to Zhang et al. (2018a) mouse ovarian granulosa cells and estradiol hormone rejuvenation including the LHR transcription were interlinked with Smad9 expression while its upregulation unveiled negative feedback on FSH secretion and granulosa cells proliferation. Another study by Diaz et al. (2011) in Gallus domesticus ovarian follicles revealed the up-regulation pattern of Smad9 mRNA transcripts in an ascending order based on size, which was approximately 10-20-fold expression difference compared to other selected candidate genes of the study. In reference to Yu et al. (2019), Smad9 gene and protein clampdown noticeably increased the annual laying performance in Chinese Wanxi white geese. This was correlated with an increased amounts of LHR expression and Estradiol biosynthesis which are the key hormones in follicle viability for ovulation while no evident shifts observed in expression levels of FSH and progesterone (P4). It was also demonstrated that the variations in Smad9 gene structure can be related to egg laying performance in Wanjiang white geese and hence reaffirmed its potential to be used as genetic marker in selective breeding of prolific geese (Xu et al., 2015).

Inhibins

Inhibins belong to the transforming growth factor-β (TGF-β) superfamily, reproductive hormones secreted mainly by testes and ovaries acting as negative modulators of FSH and characterized by their antagonistic interaction with activins (Looyenga et al., 2010). Inhibins comprise of α (INHA) and β (INHBA) subunits linked by disulfide bonds actively forming inhibin A (α-βA) and B (α-βB) respectively (Meunier et al., 1988). Inhibins have been implicated in female reproduction physiology such as oocyte and follicle maturation, and embryogenesis (Sirotkin, 2011). Previous studies revealed that INHA expression is associated with regulatory functions during folliculogenesis by controlling cell proliferation, sex hormones levels and programmed cell death in the chicken (Cui et al., 2019, 2020, 2021). The study of function has also shown that the shutdown of inhibin α subunits increased the granulosa cell death via numerous apoptotic signaling pathways and altered the hormonal production in the mice (Kadariya et al., 2015). In the bovine the in vitro supplementation of granulosa cells (GC) with inhibin A downregulated the automated cell death induced proteins and hence increasing GC viability in a dose dependent manner (Xu et al., 2020). Further study elucidated the presence of inhibin α transcripts and proteins were abundant in sterile allotriploid crucian carp ovaries and testes as compared to diploid red crucian carp (Huang et al., 2021a). According to Chen et al. (2007), inhibin α gene was dominantly expressed in healthy F1 follicle GC than in GC of follicles undergoing atresia and suppressing this gene under laboratory conditions elevated the granulosa cell death with reduced estradiol levels in Yangzhou geese. Effective immunization against inhibin improved the egg laying performance in Magang and Landaise goose breeds (Huang et al., 2007). According to Qin et al. (2013a), INHA and INHBA were highly expressed in GC of mature follicles suggesting their potential in endocrine activities associated with follicular recruitment towards dominant follicles selection and oviposition. It was further reported that Zhedong geese showed reduced brooding period intervals associated with an increasing amount of luteinizing hormone when immunized against INHA anticipating an increase on egg laying trait (Zhang et al., 2021b).

Melatonin

Melatonin (N-aceyl-5-methoxytryptamine) is a chief hormone secreted by pineal gland involved in number biophysiological activities including cytoskeletal organization, biological rhythms, antioxidative associated stress, digestion, neurons, and reproduction in animals (Ezzati et al., 2021). The production of melatonin is dependent on the day and night longevity cycle activating necessary biological processes in seasonally breeding organisms (Tabecka-Lonczynska et al., 2017). Detection of melatonin membrane receptors (MT1 and MT2) and enzymes associated with melatonin synthesis arylalkylamine N-acetyltransferase (AANAT) and hydroxyindole‐O‐methyltransferase (HIOMT) in corpus luteum in sheep (Xiao et al., 2018) and sows (Zhang et al., 2018b) indicating that melatonin may influence reproduction cycle. Melatonin action through its receptors improved the primordial follicle stimulation which involved the regulation of developmental mechanisms of follicular membranes including the sex steroids in in vitro in bovine (Feng et al., 2018b; Wang et al., 2012, 2017, 2018) and mare (Pedreros et al., 2011). In vitro studies have confirmed that melatonin can enhance the survival and approve meiotic capability of oocytes, upsurges the mitochondrial action supported by decline in ROS production nonetheless of administered dosage in sheep ovarian follicles (Barros et al., 2020a, b). Single nucleotide polymorphisms (SNPs) of high-affinity melatonin receptor subtype genes (MTR1A, MTNR1B , and MTNR1C) were quantifiable between closely related species and different organisms linked with the fecundity and seasonal breeding indicating their functional contribution in regulating reproduction traits (Feng et al., 2018a; He et al., 2019; Naby and Basha, 2016). In accordance with the previous study by Tamura et al. (2017), melatonin downregulated number of genes associated with ovarian aging in mice and also reported to accelerate puberty on set in mice by promoting FSH synthesis (Yang et al., 2021). These outcomes underline the importance of melatonin in increasing oocyte viability. In chicken, exogenous melatonin triggers mTOR signaling cascade through its receptors, a signaling pathway that promotes folliculogenesis with elevating sex hormones, antioxidant enzymes and controlling granulosa cells propagation and death (Hao et al., 2020a, b). In mares, melatonin deferred the embryonic development persuaded by the reduced levels of prolactin and inverse increase in levels of FSH and luteinizing hormone (Wang et al., 2014). According to He et al. (2014) MTNR1A, MTNR1B, and MTNR1C were predominantly expressed in preovulatory follicles with cumulative tendency from early developmental stage and declined significantly in postovulatory follicles in Sichuan white goose. Another study on identification of SNPs in the goose MTNR1A gene revealed the notable variation within the genotypes that could be associated with egg production (Alsiddig et al., 2017).

Conclusion

In conclusion, avian egg production is a complex interaction between intra- and inter- ovarian signaling pathways orchestrated by hypothalamic pituitary gonadal axis intertwining with multiple growth factors and genetic regulators influencing ovarian follicle maturation. Pertaining to shortening of seasonal brooding in geese, several technical research focused on investigating potential molecular regulators involved in folliculogenesis such as heat shock proteins, and melatonin receptor genes together with some growth factors such as AMH, inhibins, BMP4, and Smad TGF families. The findings of these studies provided insightful information towards increasing egg yield in different goose species. This review summarized the available information on the selected biological regulators intended to improve goose egg production. More studies need to be carried out in order to reveal how these biological processes affect goose laying performance in a species dependent manner and their significance in molecular breeding.

Declarations

Acknowledgement

Author are grateful to Lineo Mabesa and Rethabile Mokete for improving the language.

Funding

This work did not receive any funding.

Generative AI and AI-assisted technology statement

The authors declare that no generative AI and AI assisted technology was used in the creation of this manuscript.

Statement of conflict of interest

The authors have declared no conflict of interest.

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