Acta Veterinaria Hungarica 56 (3), pp.

369378 (2008)
DOI: 10.1556/AVet.56.2008.3.10

PHYSIOLOGICAL MECHANISMS OF OVARIAN FOLLICULAR

GROWTH IN PIGS A REVIEW
Tomasz SCHWARZ*, Marcin KOPYRA and Jacek NOWICKI
Department of Pig Breeding, University of Agriculture, Al. Mickiewicza 24/28,
30-059 Cracow, Poland
(Received 23 May 2007; accepted 7 November 2007)

Follicular growth after antrum formation is determined by folliclestimulating hormone (FSH). Only two ways are possible for recruited follicles,
continuing development or atresia. In gilts, intensive ovarian follicular growth begins between 60 and 100 days of age, and fluctuations of the ovarian morphological status last about 20 days; however, at that time there are no really large follicles. Final follicular development is under luteinising hormone (LH) control; this is
why the attainment of puberty is related to an increase in serum oestradiol to a level
that causes a preovulatory surge of this gonadotropin. The pool of follicles at the
beginning of the oestrous cycle is about 3040, most of which are small (< 3 mm)
and growing. Then, the pool of follicles increases to about 80 in the mid-luteal
phase but about 50 of them are small and 30 are medium sized (36.9 mm). Some
of these follicles are in the growing phase, but some are atretic. Between days 7
and 15 of the oestrous cycle the percentage of atretic follicles fluctuates between
12 and 73%. At that time there are no large (> 7 mm) follicles because of the suppressing effect of progesterone. The number of small follicles declines after luteolysis. From the pool of medium follicles, large follicles are selected under the influence of LH, but about 70% of the medium-sized follicles become atretic. Because of the long-lasting selection process there is a significant heterogeneity in
the diameter of large follicles in oestrus. However, the number of follicles correlates with the number of corpora lutea after ovulation. Individual follicular development and the relationship between follicles are still poorly known. The use of
ultrasonography may give a closer insight into these phenomena.
Key words: Folliculogenesis, pig, gilt, sow
*

Corresponding author; E-mail: rzschwar@cyfronet.krakow.pl; Fax: 0048 (12) 633-3307

0236-6290/$ 20.00 2008 Akadmiai Kiad, Budapest

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SCHWARZ et al.

Folliculogenesis is one of the most important physiological events significantly affecting the success of animal breeding. This is why researchers and
breeders have been interested in its regulating mechanisms as well as in the possibility of manipulating that process. The mechanisms of follicular growth were
firstly recognised in cattle, and a wave pattern theory was described (Rajakoski,
1960). The basic stages of ovarian follicular growth in cattle are recruitment, selection, dominance, and according to the oestrous cycle phase atresia or ovulation. A similar folliculogenesis model, based on the wave pattern theory, was
described for sheep and goats; however, in these species the nature of some follicular development stages is different from those in cattle. First of all, this difference concerns a significantly less pronounced dominance stage, which is directly connected with an ovulation rate higher than in cattle (Schwarz and Wierzcho, 2000; Ziba et al., 2002). Despite the anatomical and physiological differences, the nature of folliculogenesis in horses, especially as regards the regular
follicular waves, seems to be very similar to that in ruminants (Ginther, 1993).
The most evident trait of sows is their high reproductive potential related
to the short farrowing interval and the high litter size. Their productive specificity is connected with physiological mechanisms of follicular growth and development, as well as with the high ovulation rate (Knox et al., 2003).
Folliculogenesis and the mechanisms regulating it differ according to female
puberty degree and physiological status (Brssow et al., 2002; Rtky et al., 2005b).
Folliculogenesis in prepuberal gilts
The ovaries of gilts become sensitive to gonadotropins at about 60 days of
age. At this point the development of tertiary follicles is observed. Such follicles
are able to synthesise follicle-stimulating hormone (FSH) receptors, so they can respond to the natural changes in serum gonadotropin concentrations as well as to
exogenous hormonal treatment (Rtky and Brssow, 1998; Rtky et al., 2005b).
The increasing role of growing follicles in oestradiol production and secretion activates a negative feedback to the hypothalamo-pituitary-ovarian axis. This causes
a decrease in serum gonadotropin concentrations at about 100 days of age in intact
gilts in comparison with ovariectomised animals (Wise, 1982). After the 100th
day, a rise in urinary oestrogen concentration was observed, which suggests a
higher metabolic rate and thus a higher activity of oestradiol. Probably this is the
reason why the levels of this hormone remain relatively low despite the increase in
ovarian follicular activity (Camous et al., 1985). A significant increase in serum
oestradiol concentration was observed directly before the first ovulation and was
associated with the first oestrous behaviour (Esbenshade et al., 1982).
The synthesis of FSH receptors makes it possible for follicles to grow to a
size over 1 mm, but their development becomes dependent on the serum FSH concentration. The intensity of atretic changes in follicles with a diameter < 1 mm is
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low (Driancourt et al., 1995). Atresia mostly occurs when follicles are between
1 and 5 mm in diameter, after the decrease in serum FSH level (Dufour et al.,
1988). Some of the follicles > 4 mm in diameter produce luteinising hormone
(LH) receptors, which make them independent of FSH concentration. This is
why exogenous FSH administration induces intensive growth of small and medium follicles, but the reaction of large follicles is weak (Guthrie et al., 1990).
Pulsatile secretion of LH at the time of FSH decrease enables further growth of
the LH-dependent follicles, while the rest of follicles start to undergo atresia. A
high level of atretic follicles 15 mm in diameter is the indicator of follicular selection process. The stage achieved by large, selected follicles does not prevent
them from atresia; however, the intensity of that process in follicles with a diameter > 6 mm is much weaker (Prunier et al., 1993).
A 20-day cycle of ovarian status changes was evaluated by laparoscopic
diagnosis of follicular growth carried out once in every 5 days in prepuberal gilts
(Bolamba et al., 1994). Three morphological types of ovaries were distinguished:
HT honeycomb type, GT grape type, IT intermediate type. The HT ovary
represents follicular growth before selection, with great number of small follicles
and low number of medium and large follicles. During follicular selection the
number of small follicles decreases while that of large follicles increases. At that
moment the morphological status of the ovary is called IT; however, it is difficult
to recognise this stage because of its short duration. GT is the ovary after selection, with a high number of large follicles (diameter > 6 mm) and a low number
of small follicles. The duration of GT is significantly shorter than that of HT (Bolamba et al., 1994). The response to exogenous gonadotropin treatment is stronger
(higher ovulation rate) in GT than in HT ovary (Bolamba et al., 1992).
Folliculogenesis in cyclic gilts
The main change in folliculogenesis after the attainment of puberty in gilts
is the appearance of the cyclic corpus luteum, which undoubtedly influences follicular development (Murphy et al., 2001). Corpus luteum activity determines the
length of the oestrous cycle because the duration of the follicular phase is generally stable. The progesterone secretion profile is characterised by a sinusoid
curve with highest values between days 10 and 14 of the cycle and a nadir in the
periovulatory period. The relationship between the number of corpora lutea and
the concentrations of progesterone is not clear. While some authors found a positive correlation (Mariscal et al., 1998; Mao et al., 1999), others did not, or even
reported a negative correlation (Egerszegi et al., 2003; Rtky et al., 2005b).
The number of small follicles is low in the periovulatory period but it increases rapidly after ovulation. The medium follicles appear at the 3rd day, however, their number becomes significant on the 5th day (Knox et al., 2003). The percentage of atretic follicles on that day is low, because the majority of them are in a
growing phase. The number of atretic follicles rises up to 50% on day 7. Between
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SCHWARZ et al.

days 7 and 15 of the oestrous cycle the percentage of atretic follicles fluctuates between 12 and 73% (Guthrie et al., 1995). This means that follicles grow in groups
with intensification of development at the same time, and then atresia follows
steadily, suggesting a kind of regularity (Guthrie et al., 1995). Nevertheless, during
the luteal phase there is a lack of really large follicles potentially able to ovulate.
Such follicles appear only just after luteolysis, i.e. after day 15 of the cycle (Lucy
et al., 2001; Guthrie, 2005). This means that follicles are not able to pass the selection process successfully and to reach maturity in the presence of corpora lutea.
The follicular phase is much shorter than the luteal phase, but there are
much more follicular events during this period. The majority of follicles of all
diameter classes are healthy and grow at the time of luteolysis. At about the third
day of the follicular phase, many follicles, especially those of small diameter,
become atretic as a result of the selection process. On the 5th day after luteolysis
all small follicles and approximately 70% of medium ones decrease in their diameter, and most of the ovarian surface is covered by large, oestradiol-active follicles in the preovulatory stage (Guthrie, 2005).
The most important event during the recruitment process is the synthesis
of FSH receptors by small follicles, which enables the latter to grow over 1 mm
in diameter (Guthrie et al., 1995). During the periovulatory period a surge of
FSH is observed, and elevated concentrations of that hormone are maintained until about day 4 or 5 of the oestrous cycle, stimulating the development of small
and medium follicles and reducing atresia (Knox et al., 2003) (Fig. 1A, B). FSH
secretion is under hypothalamic GnRH control. However, regulating the release
of pituitary hormones is not the only role of GnRH, because in gilts immunised
against GnRH the follicles are not able to grow over 1 mm in diameter, even after PMSG administration (Esbenshade, 1987). The concentration of FSH decreases at about the 5th day, and at the same time an elevated level of follicular
atresia is observed (Guthrie et al., 1995) (Fig. 1D, E). Later, in the period of high
progesterone concentration, only slight fluctuations in FSH level are observed
(Knox et al., 2003). On day 14 or 15 luteolysis occurs and the level of progesterone decreases. At that time a second significant increase in FSH concentration is
observed, which is pivotal for the recruitment process (Knox et al., 2003)
(Fig. 1G, H). Higher level of FSH was observed in gilts with a high ovulation
rate than in those with a low ovulation rate (Knox et al., 2003; Guthrie, 2005),
but some authors did not find such a correlation (Hunter et al., 1993; Mariscal et
al., 1998). The number of growing follicles in the early follicular phase is lower
than the ovulation rate. This means that in pigs, as opposed to ruminants, follicular recruitment is a long-lasting process. At the beginning of the selection process follicles from the recruited pool are in different stages of development. This
is why even selected follicles show a large heterogeneity in diameter, morphology and hormonal status, and follicular response to LH surge may be different
(Hunter et al., 1989; Hunter and Wiesak, 1990).
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LH is much more versatile in activity than FSH, because it influences not

only follicular growth but also final oocyte maturation, ovulation, and corpus luteum formation (Fig. 1N). The level of LH during the luteal phase and at the beginning of the follicular phase is relatively stable (Knox et al., 2003); however,
the phenomenon of pulsatile secretion is observed during follicular selection
(Driancourt et al., 1995) (Fig. 1J). The mechanism and duration of the selection
process in pigs are not fully recognised yet. The basis of this process is the synthesis of LH receptors in follicles (Fig. 1K). This is why the growth of such follicles becomes independent of FSH concentration. The above-mentioned pulsatile
LH secretion enables further follicular development despite the low basic level of
this hormone. The decreasing FSH concentration causes atretic changes in follicles that did not synthesise LH receptors (Hunter and Wiesak, 1990; Prunier et
al., 1993). The indicator of follicular maturity is activation of the enzymatic
aromatase complex in the granulosa cells, which enables the synthesis and secretion of oestradiol and inhibin (Yen et al., 2005). The concentrations of these hormones are stabilised on a low level during the luteal phase. After luteolysis, a
gradual increase in the concentrations of oestradiol and inhibin is observed, and
the peak occurs in the periovulatory period, enabling the manifestation of oestrus
and provoking the LH surge (Hunter et al., 1989; Knox et al., 2003) (Fig. 1L, M).
The level of both hormones rapidly decreases before ovulation, because the final
maturation of follicles is connected with a reduction in oestradiol production and
an increase in progesterone synthesis (Brssow et al., 2002; Rtky et al., 2005a).
This profile of follicular hormones suggests that there is only one selection process during the whole oestrous cycle, and the factor required for its occurrence is
low progesterone level.
Like recruitment, the process of follicular selection seems to last longer in
gilts than in ruminants, and therefore a larger number of follicles can successfully pass it. The mechanism determining the duration of selection remains unknown. Nevertheless, early studies investigating this problem showed that the
number of mature follicles and the concentration of oestrogens have a significant
role (Coleman et al., 1984). However, exogenous gonadotropin treatment causes
a superovulatory effect only if it is administered before the 18th day of the oestrous cycle (Hunter et al., 1976).
An increase in ovulation rate can be reached by exogenous hormone treatment in two ways. The administration of FSH or eCG in the period around
luteolysis increases follicular recruitment and reduces apoptosis, so more follicles can avoid atresia and continue to grow in the follicular phase (Knox and
Zimmerman, 1993; Liu et al., 2003). In turn, the purpose of LH or hCG use is not
to increase the number of small follicles in recruitment but to make it possible for a
higher number of follicles to pass selection process (Guthrie et al., 1990; Holtz
and Schlieper, 1991). The increase in ovulation rate after gonadotropin administration is more evident in monoovular species such as ruminants. Nevertheless,
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SCHWARZ et al.

GnRH

(-)

(-)/(+)

hypotalamus
hypothalamus

(-)
pituitary

(-)/(+)

(+)

(+)/(+)

(+)
(+)

A
5

(+)

4,5

Pulsatile
secretion of LH

(-)

1 2 3 4 5 6 7
hours

(+)

(+)

3,5

3
2,5

FSH

(-)

LH

1,5
1
0,5
0

FSHR

(+)

E
FSHR

10 11 12 13 14 15 16 17 18 19 20 21

(-)

H (+)
FSHR

FSHR

FSHR

K
FSHR

FSHR

(+)

LHR

LHR

N
(+)

CL

CA

luteal
phase

10
9

follicular
phase

(-)

ovulation

(+)

(+)

CB

F L

H (+)

5
4
3
2

(+) recruitment

atresia

recruitment

selection

atresia

10 11 12 13 14 15 16 17 18 19 20 21

70

60

50

progesteron
progesterone

40

30

20

oestradiol
inhibin
estradioland
i inhibina

10

10 11 12 13 14 15 16 17 18 19 20 21

daysofofthe
theoestrous
estrous cycle
Days

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375

problems occur after ovulation because of the formation of ovarian cysts

(Schwarz et al., 2007a; Schwarz et al., 2007b). In gonadotropin-treated gilts, the
higher ovulation rate does not guarantee an equal increase in the number of piglets born. This is why superovulation is not a widespread method in pig breeding
(Holtz and Schlieper, 1991). The synchronisation of ovulation is more important,
especially in large farms, and the use of GnRH agonist inducing an endogenous
LH release seems to be the most effective for this purpose (Brssow et al., 2007).
Folliculogenesis in multiparous sows
The reproduction process of multiparous sows is characterised by the lack
of a luteal phase. The farrowing interval consists of gestation, lactation, and a
short weaning to oestrus interval lasting 47 days, which includes only the follicular phase finished by ovulation. After successful mating or insemination, the
subsequent gestation and farrowing interval begins.
During lactation, follicular development is limited and there is no selection process at all because of the low serum gonadotropin, especially LH, concentrations. This is why the occurrence of oestrus and ovulation is impossible
during that period (De Rensis and Foxcroft, 1999; Mao et al., 1999). Weaning is
associated with strong, acute stress, even if it is a single and short event. The sow
responds with an activation of the hypothalamo-pituitary-adrenal axis and an increase in the secretion of cortisol, which enables the restoration of homeostasis.
Then the concentration of cortisol decreases, and the high level of FSH enables
recruitment from the follicular pool. The number of follicles in the cohort depends on the FSH level (Wheaton et al., 1998). However, the most important factor is the pulsatile LH release, which enables follicles to pass the selection process and to begin oestradiol synthesis and secretion (Tsuma et al., 1995). The frequency of LH pulses significantly influences the weaning to oestrus interval (van
den Brand et al., 2000). Excessive stress at the time of weaning connected, e.g.
with dominance-hierarchy fights results in the maintenance of high cortisol level
and the reduction of pulsatile LH release (Tsuma et al., 1995). In this case, the
disruption of follicular development may be the cause of pathological anoestrus

Fig. 1. Follicular development during the oestrous cycle in gilts. FSH release at the beginning of
the oestrous cycle (A) stimulates the growth of small and medium follicles which synthesise FSH
receptors (FSHR) (B). Increasing progesterone level prevents the LH surge (C) and decreases FSH
concentration (D), which causes follicular atresia (E). Decreasing progesterone level after luteolysis (F) enables an increase in FSH concentration (G) and another follicular recruitment (H), as well
as the pulsatile secretion of LH (J), follicular selection and maturation of follicles with LH receptors (LHR) (K). Increasing oestradiol concentration connected with follicular development after selection (L) decreases FSH level (I), causing atresia of follicle FSHR. However, high oestradiol
concentration and low progesterone level enables the LH surge (M), causing ovulation of large follicles (N) and the formation of corpora lutea, which initiate another oestrous cycle (based on Driancourt et al., 1995; Guthrie et al., 1995; Lucy et al., 2001; Knox et al., 2003; Guthrie et al., 2005)
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SCHWARZ et al.

after weaning (van de Wiel and Booman, 1993). The morphology of follicles as
well as the processes which regulate folliculogenesis during recruitment and selection are identical with those in the follicular phase of the oestrous cycle in cyclic gilts, described previously. However, the time necessary for follicles to
achieve preovulatory maturation seems to be longer, which may explain the longer
oestrous cycle of sows as compared to gilts (Rtky et al., 2005a).
There are still many unclear aspects of follicular development during the
oestrous cycle in pigs, especially in the luteal phase. Waves of follicular development were not observed in gilts and sows; however, in small ruminants the
wave pattern theory was controversial until ultrasonography was introduced into
the daily examinations of ovaries. Most studies on porcine reproduction employ
repeated endoscopic observations. Compared with ultrasound imaging, an advantage of this method is that the quality of ovarian features can be evaluated (Rtky
et al., 1998). However, it is impossible to use this method for daily observations
over a long period of time necessary for evaluating the pattern of individual follicular growth and the relationship among follicles day by day. Meanwhile, a better recognition of the relations mentioned above may help in the improvement of
reproductive performance parameters in the breeding practice and in working out
methods for the artificial control of the oestrous cycle and ovulation rate.

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Acta Veterinaria Hungarica 56, 2008