Anime Ova

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anime ova

Follicular wave of Mammals A Review and Web Briefs

Follicular phase of menstrual cycle

OF menstruation in women's follicular Artesia is completed. All of developing follicles die down, only one pre-ovulatory follicle remains. The process of apoptosis kills or destroys all other follicles in their oocyte inside. The causes of Artesia is unknown, but FSH helps to prevent it. The recent investigations of the real factors that have to oocyte well done this, ovulate, it was reported. [20 manner manner and

INTRODUCTION

Follicle development resembles the cattle, humans and equines. Bovines are more similar of people than equines for their cycle length and poly oestrus behavior (Baerwald, 2009)

Follicular development is reported in the waves. The waves were preceded by changes in hormonal status ie estrogen, progesterone and FSH and LH ratio balances. Selection of a dominant follicle can occur in each wave.

The estradiol, inhibin A, and IGF-II act

is giving way to dominant follicle to grow at a

decreasing FSH and LH increasing environment. The subordinate follicles Artesia and tasted death.

GAMETES history

Hippocrates in the 5th century BC

Did male and female menstrual blood ejaculate join to form the new born.

Aristotle knew that women contributed to the baby thing and men gave vent to form and shape in the embryo.

Accurate reference development of humans and swivel from moving water or the sperms are described in the book of GOD as revealed to Mohammad Hazrate Wahi (trust) in 550 AD. So the Quran is the first written reference to the procreation of gametes and a mechanism of its development in the uterus too is narrated. The excitement and a fixed rate of reduction of the pregnant uterus described the Quran is nevertheless fortified with scientific investigations. So Quran is the first written reference to fetal development in the world. Manner, 2009,2010)

William Harvey, Ex postulation OVA OMENA mean everything comes to eggs in the 17th century AD was confirmed by

Rangier de Graaf in 1672.He mistook the follicle as an egg and it's still named after him as Graafian follicle.

It is only in the 19th century, von Baer found the egg follicle and

ovarian follicle and folliculogenesis is recognized, (manner, 1984, Cobb, 2006)

That was in the 20th century,

The human egg was first discovered at the

Antral follicular growth is reported. (Cobb, 2006).

Many specific details of folliculogenesis sa mammals will be studied and be in-vitro experiments enhanced our understanding processes involved in different species (Table 1]

Table 1-History of folliculogenesis.

Author year species

Bullough Mandle and Zuckerman 1946 Mice Mouse 1950

1951 block Green and Zuckerman 1951 Female Monkey

Maybe Rajakoski 1960

Brand and deJong 1973 sheep

Clark et al 1975 Pigs

1976 Goats manner

1984 Sheep manner

Ginther and Pierson, 1984 horse

Adams et al 1989 came

(McCorkell et al 2006 wapiti

Asher et al 1997 Deer

Figure 1.Folliculogenesis website diagram.

The diagram shown below taken from a free website shows the development

process of follicle from the bud epithelium, but a new concept discussed later in this book should evidences indicate that the surface epithelium differentiate too oocytes and is surrounded of the follicular wall and a number of videos on the web provide figure animation and audio commentaries

 

Using by ultrasonography was instrumental in the study [Ginther et al, 2004] The transition from m ovarian reserve in growing follicles is a continuous process in women. Fig 2 & 3.below.

Fig 2

Figure 3

TIME THE sequence of follicular wave

Antral follicles seen the young 14 days old

[Evans et al, 1994]

Prenatal follicles might nonresponsive sa gonadotrophins but early antral follicle I respond to gonadotrophins [Adashi, 1997] In the women starting from the follicle is recruited and destines to develop in a sequential manner throughout reproductive life. [Baird, 1987] The recruitment of antral follicles are produced both in early and late luteal phases of the cycle. [McGee and Hsueh, 2000] The recruitment of 2-5 mm follicles is an ongoing process after puberty. [Gougeon, 1979] A theory suggests that the follicles are recruited in response to the increase in GnRH and FSH lutuem regresses once the corpus [Hodgen, 1982] The follicular phase follicles are 2mm in diameter. Their growth is confusing under the influence of estrogen, progesterone, FSH AND LH [ET AL Baird, 1975] These concepts are changing and update the latest published as a review. [AT manner manner 2009.2010] In bovines and equines the ovulatory follicles were seen during proestrus, oestrus saw last maturation and ovulation, Corpus lustrum develops during oestrus and met diestrum when CL is functional and secretes progesterone. The oestrus cycle of cows is 21 days, consisting of 4 days follicular luteal phase and rest as a part, etc. have 7 days long follicular phase and luteal phase. Women have a 14 day follicular phase and nearly half of the full cycle of 28 days. a

Decline in circulating FSH and increase in follicular

estradiol, inhibin A, and IGF-II to collective action

dominant follicle to enable continue to grow at a

endocrine environment of decreasing and FSH

LH increase, while the underlying follicles undergo

return.

Chapter 3

.

Initiation of follicle development

The Journey

preovulatory follicles is estimated that more than 175 days in women.160 day cattle, [Gougeon, 1986] A second follicle to ovulatory follicle takes about 42 days. Research in this aspect has been reviewed and published. [Bearwald 2009]

Baerwald. Comparative folliculogenesis.

Six. Reprod., 22 v.6, n.1, p.20-29, January / March 2009 refers

Many author has named the developing follicle as dominant, privileged, Doubtful and atreteic follicles as complementary or sub dominate follicles. [BAERWALD, 2009)

Deviation from the dominant follicle

largest subordinate follicle occurs at a diameter of 9 mm

sa heifers, 10 mm in women and 23 mm sa mares

(Gastal et al., 1997, Ginther et al., 2001; Baerwald et

al., 2003).

Alignment with ovulatory wave occurs, the

average, 4 days after the emergence of the largest follicle to

13 mm in mares and 6 mm in women (Ginther et al.,

2004), compared to 3 days after the emergence of

largest follicle at 4 mm in cattle (Ginther et al., 1997).

The dominant follicle maintains a constant growth rate

deviation throughout the process, while the assistant

follicles exhibit a reduction in growth rate (Gastal et

al., 1997, Ginther et al., 2001a). No evidence to

women, mares and heifers, the dominant follicle

exhibits an early size advantage over the other follicles in

the pool, enabling is to establish dominance before

subordinate follicles reach a similar diameter (Ginther et

al., 2001a; Ginther et al., 2004).

Baerwald. Comparative folliculogenesis.

Six. Reprod., 24 v.6, n.1, p.20-29, January / March 2009

Figure 2. Morphologic and endocrinologic characteristics of ovarian follicular wave dynamics in women. Follicle

and luteal dynamics is illustrated in women with 2 (A) and 3 (B) follicle waves during an IOI. Follicles outlined

a dashed line represents the large waves that were present in some, but not all women. Serum concentrations of

Estradiol and progesterone in women with 2 and 3 curves are shown (C). Serum concentrations of FSH and LH sa

women with 2 against 3 follicle waves during an IOI is shown (D).

A

B

C

D

Baerwald. Comparative folliculogenesis.

Six. Reprod., V.6, n.1, p.20-29, January / March 2009 25

Figure 3. Profile of the individual identified in the follicles for 3 horse mares during the oestrous cycle, starting from day

the first ovulation and ends 4 days after the second ovulation (ov). Follicles smaller than 10 mm were removed. A

anovulatory dominant follicle of a large wave (MW) and a minor wave (MW) is shown followed by the ovulatory

wave of mares 1 and 2, respectively. Mare 3 had no significant follicles grow during the first days after

ovulation. (Figure courtesy of EL Gastal).

Figure 4. Schematic of two-follicle model illustrating the size advantage of the future dominant follicle. Based on

limited information, the diameter scale for women is speculated to be similar scale for heifers. The extent and

amount of parallelism between the two follicles within developing common form varies great

individual and exaggerated in the illustration. On average, the common-growth phase ends and begins diverting

when the largest follicle reached sa indicated diameters. Alignment is established before the next largest follicle can

reach a similar diameter, represented by the width of the vertical bar. (Fake with permission, Ginther OJ.

2001. Biol Reprod, 65:638-647).

Physiologic mechanism underlying follicle

dominance

Physiologic selection of a dominant follicle is a

complex phenomenon, which is regulated by of endocrine,

autocrine and paracrine factors. Most research

accurately characterize the underlying approach

selection so far was performed on animals.

Women obtained results appear consistent with the

domestic farm animals.

The increase in FSH responsible for stimulating

follicle recruitment begins to decline with

selection of dominant follicle and the Artesia

subordinates (Santbrink et al., 1995; Gastal et al., 1997;

Ginther et al., 1997). The duration and size

FSH rise above a critical threshold is shown in

determine the number of follicles selected from

recruited group (Brown, 1978; Baird, 1987, and Fauser

Heusden, 1997). Both FSH, heifers and mares

exhibit a small but significant transient increase in

circulating around of LH during the deviation (Ginther et

al., 1998, 2001c). A temporal increase associated LH

follicle the selection of the women are not seen.

The dominant follicle exerts the same morphologic

and performance of dominance over other follicles of

wave. Circulating concentrations of estradiol plus

the continued growth of the dominant follicle to

women, mares and cattle (McNatty, 1981: Baird, 1983;

Baerwald. Comparative folliculogenesis.

Six. Reprod., 26 v.6, n.1, p.20-29, January / March 2009

Santbrink et al., 1995; Gastal et al., 1999, Ginther et al.,

2000a). The dominant follicular fluid of the follicles

contains more women estradiol and progesterone

lower levels and androstenedione levels than

subordinate follicles (McNatty, 1981; Schneyer et al.,

2000), consistent with the findings in mares (Donadeu and

Ginther, 2002). Sa heifers, however, estradiol and

androgen concentrations increase in the development of

dominant follicle (Ask et al., 2002). Dominant follicle

estradiol production is believed to provide negative

feedback on FSH and induce the formation of granulosa

Cell LH receptors, which initiates a transfer from sa FSH

LH dependency on dominant follicle (Yamoto et al.,

1992b, Xu et al., 1995, Bodensteiner et al., 1996; Gastal

et al., 1999, 2000, Sullivan et al., 1999, Ginther et al.,

2001b, c). The dominant follicle then becomes unique to the

its ability to thrive despite decreasing FSH, while

subordinate follicles regress.

Although the dominant follicle performing the basic

paper, all the follicles of an emerging wave of contributions

suppression of wave-eliciting roll in FSH in cattle

(Ginther et al., 2000b). Recruited follicles within

inhibin group produces additional activities to curb

FSH in women, cows and mares (Ginther et al., 2001a).

The selection process in women is accompanied by a

lessening sa circulating inhibin B and inhibin increase

The concentrations (Yamoto et al., 1992a, Roberts et al.,

1993; Schneyer et al., 2000). Distinct functions of an inhibin

and inhibin B, however, during the conflict with follicle

equine and maybe oestrous cycle has not been shown

(Ask for alms and Ginther, 2006). The role of activin and

follistatin in regulating follicle selecting people

(Roberts et al., 1993; Schneyer et al., 2000) and

domestic farm animals (Donadeu and Ginther, 2002;

Glister et al., 2006) have been evaluated. However,

results were inconclusive and further investigations are

required. An increase in free insulin-like growth

Factor (IGF) in follicular fluid of dominant

follicle, mediated by IGF binding proteins proteases -4/-5

(The equivalent of Pregnancy Associated Plasma ox

Protein-A, PAPP-A) has also been implicated as a

Candidates for adding responses to the Ability

gonadotropins and thereby the onset of follicle choice

cattle and horses (for reviews, see Fortune et al., 2004;

Beg and Ginther, 2006). Similarly, studies in women

have reported an increase in IGF-II and IGFBP-4

protease (PAPP-A) along with follicle selection

(Review sa Guidice, 1995).

Preovulatory follicular growth

The dominant follicle grows at a rate of

approximately 1.2 mm / day in cattle, 2.7 mm / day

mares, and 1.8 mm / day in Following its women

selection until it ovulates mid-cycle (Ginther et al.,

1989a; Gastal et al., 1997, Ginther et al., 2004). The

percentage of diameter increase was similar between

species, given the size difference leading

follicle to alignment. The dominant follicle in women

ovulates a diameter of approximately 20 mm (Pache

et al., 1990, Baerwald et al., 2003a). In contrast,

preovulatory diameter of dominant follicle to

might be smaller (16 mm) while the ass is

big bigger (45 mm).

Ovulatory dominant follicle growth

results in a rapid elevation of circulating estradiol

cattle, mares and women (McNatty, 1982, Sunderland et

al., 1994; Gastal et al., 1999). Estradiol production from

The dominant follicle peaks a day before the LH surge

among women, three days before the LH surge sa mares, and

on the day of LH surge in cattle. Dominant follicle

estradiol provides positive feedback

hypothalamus and pituitary to stimulate the release of

LH required for inducing ovulation. The estradiol

levels in mid-late follicular phase increase earlier in

women with 2 versus 3 follicular waves, and

preovulatory estradiol peak occurs 2 days earlier

women with two waves (Baerwald et al., 2003a).

Similarly, the preovulatory FSH and LH surges occur 1

days earlier in women with 2 versus 3 follicle to wave

with a shorter cycle length (Baerwald et al.,

2003a), similar to previous studies in cattle (Adams,

1999). As LH levels increase in late follicular phase,

preovulatory follicle in all three species shifts from a

estrogens-secreting state with a progesterone secreting

state and transformation from follicular cells to luteal

Cells begin.

Attribution and future directions

Waves of antral follicular development

are well-documented in several animal species,

with mares and cattle (Ginther, 1993: Fortune,

1994: Adams, 1999). Recent evidence supports

notion of antral follicular wave patterns of recruitment

in women (Baerwald et al., 2003a, b, 2005). Pattern

follicular wave emergence in women closely resemble

to the previously described during the cow and horse

oestrous cycle. The number of waves observed depending

the length of the cycle. Furthermore, the final

IOI is ovulatory wave, while all preceding waves

were anovulatory. Major and minor patterns of follicle

wave of the women are similar to those observed

equine oestrous cycle. The mechanism underlying

deviation of the dominant follicle from subordinate

follicles were similar in all three species. Thus, both

cow and horse species were established as

model for the study of human ovarian function (Adams

and Pierson, 1995, Ginther et al., 2004). Animal models

is critical for increasing our understanding of

biological mechanisms underlying ovarian

folliculogenesis, given the practical and ethical

boundaries in the study of human reproductive tissues. The

purpose of building model animals for studying human

ovarian function is to provide the basis for hypothetical

ongoing research on women, which is quite that

domestic farm animals (Donadeu and Ginther, 2002;

Glister et al., 2006) are reviewed. However,

results were inconclusive and further investigations are

required. An increase in free Insulin-like growth

Factor (IGF) sa follicular fluid of dominant

follicle, mediated by the IGF binding protein proteases -4/-5

(The cow equivalent of Pregnancy Associated Plasma

Protein-A, PAPP-A) has also been implicated as a

candidate for adding Ability to respond to the

gonadotropins and thereby the onset of follicle choice

cattle and horses (for reviews, see Fortune et al., 2004;

Beg and Ginther, 2006). Similarly, studies in women

have reported an increase in IGF-II and IGFBP-4

protease (PAPP-A) along with follicle selection

(Review sa Guidice, 1995).

Preovulatory follicular growth

The dominant follicle grows at a rate of

almost 1.2 mm / day in cattle, 2.7 mm / day

mares, and 1.8 mm / day following its women

selection until it ovulates in the middle of-cycle (Ginther et al.,

1989a; Gastal et al., 1997, Ginther et al., 2004). The

percent diameter increase was similar between

species, given differences in the size of the leading

follicle to alignment. The dominant follicle in women

ovulates a diameter of approximately 20 mm (Pache

et al., 1990, Baerwald et al., 2003a). In contrast, the

preovulatory diameter of dominant follicle to

might be smaller (16 mm) while the ass is

big bigger (45 mm).

Ovulatory dominant follicle growth

results in a rapid elevation of circulating estradiol

cattle, mares and women (McNatty, 1982; Sunderland et

al., 1994; Gastal et al., 1999). Estradiol production from

The dominant follicle Peaks a day before the LH surge

among women, three days before the LH surge sa mares, and

on the day of LH surge in cattle. Dominant follicle

estradiol provides positive feedback

hypothalamus and pituitary to stimulate the release of

LH required for inducing ovulation. The estradiol

levels in mid-late follicular phase increase earlier in

women with 2 versus 3 follicular wave, and

preovulatory estradiol peak occurs 2 days earlier

women with two waves (Baerwald et al., 2003a).

Similarly, the preovulatory FSH and LH surges occur 1

days earlier in women with two versus three follicle waves

with a shorter cycle length (Baerwald et al.,

2003a), similar to previous studies in cattle (Adams,

1999). As LH levels increase in late follicular phase,

preovulatory follicle to all three species shifts from a

state-estrogens secreting a progesterone secreting

state and transformation from follicular luteal cells

Cells begin.

Conclusions and future directions

Waves of antral follicular development

are well-documented in several animal species,

with mares and cattle (Ginther, 1993: Fortune,

1994: Adams, 1999). Recent evidence support

notion of antral follicular wave patterns of recruitment

in women (Baerwald et al., 2003a, b, 2005). Pattern

follicular wave emergence in women closely resemble

to the previously described during the cow and horse

oestrous cycle. As follow the waves depends

the length of the cycle. Furthermore, the final

IOI is ovulatory wave, while all preceding waves

were anovulatory. Major and minor patterns of follicle

wave of the women are similar to those observed

equine oestrous cycle. The mechanism underlying

deviation of the dominant follicle from subordinate

follicles were similar in all three species. Thus, both

cow and horse species established as

model for the study of human ovarian function (Adams

and Pierson, 1995, Ginther et al., 2004). Animal models

is critical for increasing our understanding of

biological mechanisms underlying ovarian

folliculogenesis, given the practical and ethical

boundaries in the study of human Reproductive tissues. The

purpose of developing animal models for studying human

ovarian function is to provide hypothetical basis for

continued research on the women, which is fully

Baerwald. Comparative folliculogenesis.

Six. Reprod., V.6, n.1, p.20-29, January / March 2009 27

lead to the development of safer and more effective

infertility and contraceptive therapies. It is further

anticipated that future research on women is caused

views on female reproductive function in animals

species.

Future studies should be performed

determine the role of the CL in regulating the fate of

follicular waves in women and domestic animals. A

greater understanding of the roles of paracrine and

autocrine factor in regulating ovarian follicular wave

is required. Repeatability of follicle pattern wave has

recently been documented in cattle during oestrous

cycle (Jaiswal et al., 2005) and some follicles

endpoints of the horse (Jacob et al., 2008). At present,

We are conducting studies in our laboratory to learn

repeatability the follicle wave dynamics in women. The

cow and horse species have recently been

established as models for studying reproductive sa roundup

women (Malhi et al., 2005, 2006, 2007, Carnevale,

2008; Ginther et al., 2008a, b). Continued to study

areas may provide insight into the age-related changes

human female reproductive potential as well as

Infertility associated with premature ovarian failure.

Acknowledgments

The author sincerely wishes to thank Dr.

Roger Pierson at the Department of obstetrics,

Gynaecology and Reproductive Sciences and Dr. Gregg

Adams at the Department of Veterinary Biomedical

Sciences at the University of Saskatchewan for their

assistance in writing this manuscript.

References

Adams GP, PG Griffin and OJ Ginther. 1989. In Situ

morphologic dynamics of ovaries, uterus, and cervix sa

Llamas. Biol Reprod, 41:551-558.

Adams GP, Matteri RL, Kastelic JP, Ko JCH,

Ginther OJ. 1992. Association between of surges

follicle-stimulating hormone and the emergence of

follicular waves in heifers. J Reprod Fertil, 94:177-188.

Adams GP, Pierson RA. 1995. Cow model for learning

of ovarian follicular dynamics in humans.

Theriogenology, 43:113-120.

Adams GP. 1999. Comparative patterns of follicle

developing and selecting ruminants. J Reprod Fertil

Suppl, 54:17-32.

Adashi ey. 1994. Endocrinology of the ovary. Buzz

Reprod, 9:815-827.

Asher GW, IC Scott, KT O'Neill, JF Smith, Inskeep

EK, Townsend EC. 1997. Ultrasonographic monitoring

of antral follicle development in red deer (Cervus

elaphus). J Reprod Fertil, 111:91-99.

Baerwald A, G Adams, R Pierson. 2003a.

Characteristics of ovarian follicular wave dynamics

women. Biol Reprod, 69:1023-1031.

Baerwald A, Adams G, Pierson R. 2003b. A new

model for ovarian follicular development during

human menstrual cycle. Fertil Steril, 80:116-122.

Baerwald AR, Adams GP, Pierson RA. 2005. Form

and corpus luteum function during the human

menstrual cycle. Ultrasound Obstet Gynecol, 25:498 -

507.

Baird D. 1987. A model for follicular selection and

ovulation: lessons from super ovulation. J Steroid

Biochem, 27:15-23.

Baird D, Fraser IS. 1975. Concentration of oestrone

sa follicular fluid and oestradiol and ovarian venous

blood of women. Clin Endocrinol, 4:259-266.

Baird DT, Baker TG, McNatty KP, Neal P. 1975.

Relationship between corpus luteum secretion of

and the length of the follicular phase of ovarian

cycle. J Reprod Fertil, 45:611-619.

Baird DT. 1983. Factors regulating growth

preovulatory follicle in sheep and man. J Reprod

Fertil, 69:343-352.

Beg MA, Bergfelt DR, intimidation K, Ginther OJ. 2002.

Follicle selection in cattle: dynamics of follicular fluid

factors during development follicle dominance. Biol

Reprod, 66:120-126.

Beg MA, Ginther OJ. 2006. Follicle selection in cattle

and horses: role of intrafollicular factors. Making a copy,

132:365-377.

Bergfelt DR, Ginther OJ. 1992. Relationship between

circulating concentrations of FSH and follicular waves

during early pregnancy mares. J Sci horse VET,

12:274-279.

Bergfelt DR, Ginther OJ. 1993. Relationship between

FSH surges and follicular waves during the estrous

cycle in mares. Theriogenology, 39:781-796.

Block E. 1951. Quantitative morphological

investigations of the follicular system in women;

variations in different phases of sexual cycle.

Acta Endocrinol, 8:33-54.

Bodensteiner KJ, Wiltbank MC, Bergfelt DR,

Ginther OJ. 1996. Alterations in follicular estradiol

and gonadotropin receptors during development

maybe antral follicles. Theriogenology, 45:499-512.

Brand A, de Jong WH. 1973. Qualitative and

quantitative micromorphological investigations of

tertiary follicle population during the oestrous cycle

sheep. J Reprod Fertil, 33:431-439.

Brown JB. 1978. Pituitary control of ovarian function:

concepts derived from gonadotrophin therapy. Austr NZ

J Obstet Gynaecol, 18:47-54.

W. Bullough 1946. Mitotic activity among women, adults

mouse. Musculus L moose, 231:453-516.

Carnevale em. 2008. The disturbance model for follicular

maturation and reproductive aging in the female.

Theriogenology, 69:23-30.

Check J, Dietterich C, Houck Tues 1991. Ipsilateral

against contra lateral ovary selection of dominant follicle

succeeding cycle. Obstet Gynecol, 77:247-249.

Clark JR, First NL, Chapman AB, Casida LE. 1975.

Ovarian follicular development during the estro

About the Author

PROF G.M.WANI(Ghulam Mohyuddin Wani )Ph.D (Animal Reproduction / Gynaecology)  IVRI, Dr. Med. Vet (Animal Reproduction/ )HUSBANDRY HANNOVER)after retirement has permanantly wrting webarticles,His articles are on free websites and have been used by more than 10,000 students in the past one year.We request the viewers of these articles to comment on the shortcomings.If no comment is offered we feel the attempt is futile.FOr our readers we keep articles without spelling check to judge their sense of peer review and correction

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