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Theriogenology 71 (2009) 1343–1357

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Review
Compilation of classical and contemporary terminology used to
describe morphological aspects of ovarian dynamics in cattle
A.T. Peter a,*, H. Levine b, M. Drost c, D.R. Bergfelt d
a
Department of Veterinary Clinical Sciences, School of Veterinary Medicine, Purdue University, West Lafayette, IN 47907, USA
b
Tufts Ambulatory Service, Department of Environmental and Population Health, Cummings School of Veterinary Medicine,
Tufts University, South Woodstock, CT 06267, USA
c
Department of Large Animal Clinical Sciences, University of Florida, Gainesville, FL 32605, USA
d
School of Veterinary Medicine, University of Wisconsin, Madison, WI 53706, USA
Received 7 August 2008; accepted 20 December 2008

Abstract
Veterinarians and scientists involved in applied and basic research in cattle require a lexicon of terms that is used uniformly so
that diagnoses and inference of results between and among studies can be correctly interpreted and substantiated or negated and
therapy and hypotheses can be formulated without unnecessary confusion and redundancy in treatments and experiments. This
review provides a compilation of many of the classical and contemporary terms used in association with ovarian dynamics primarily
during the estrous cycle in cattle, which can also apply to other reproductive states. While many classical terms used to describe
healthy and diseased conditions associated with follicles and corpora lutea are still applicable today, there are some that have
become antiquated (e.g., cystic corpus luteum, cystic ovarian degeneration, luteolysis, and granulosa cell tumor), due, in part, to
advanced technology (e.g., ultrasonography) and a more thorough understanding of ovarian function. In this regard, older terms
have been revised (e.g., corpus luteum with a cavity, follicular and luteinized-follicular cysts, structural and functional luteal
regression, and granulosa-theca cell tumor) and newer terms have been coined (e.g., follicle deviation) and advocated herein.
Defining and adopting terminology used in bovine reproduction that is clear, precise and understandable and available in a single
source, is expected to make the exchange of clinical and research information and outcomes more effective, safe, and economical.
Published by Elsevier Inc.

Keywords: Cattle; Terminology; Ovarian dynamics; Ovarian pathology

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1344
2. Estrous cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1345
2.1. Follicular phase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1346
2.1.1. Proestrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1346
2.1.2. Estrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1346
2.2. Ovulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1347

* Corresponding author. Tel.: +1 765 494 5808; fax: +1 765 496 1108.
E-mail address: petera@purdue.edu (A.T. Peter).

0093-691X/$ – see front matter. Published by Elsevier Inc.

doi:10.1016/j.theriogenology.2008.12.026
1344 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357

2.3. Luteal phase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1348

2.3.1. Metestrus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1348
2.3.2. Diestrus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1348
3. Follicular wave dynamics during the estrous cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1349
3.1. Follicular wave emergence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1350
3.2. Follicle selection and deviation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1350
3.3. Follicle dominance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1351
4. Classification of anestrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1351
5. Ovarian diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1353
5.1. Follicular and luteinized-follicular cysts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1353
5.2. Parovarian cysts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1353
5.3. Fibrin tags . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1353
5.4. Granulosa-theca cell tumor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1354
5.5. Rete ovarii (ovarian cyst) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1354
5.6. Ovarian bursa adhesions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1354
5.7. Ovarian hypoplasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1354
6. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1354
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1355
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1355

1. Introduction as well as prognosis or effectiveness of corrective and

Historically, terminology used to describe the
reproductive tract in female cattle was based, in part,
on gross examination in situ or ex situ during surgery or
necropsy, respectively. In the 1980s, with the advent of
ultrasonography and the ability to view structural
changes of the bovine reproductive organs in a
noninvasive manner over time [1–3], some of the
original terms used to describe the reproductive tract
required qualification or replacement and, in some
instances, new terms were coined and advocated.
However, modifying or replacing traditional or con-
ventional terminology used in bovine reproduction with
contemporary language is not always universally
accepted; therefore, the scientific and lay literature
and reference texts are filled with terms, definitions and
colloquialisms that are often antiquated and ambiguous.
Although reproductive disorders can be diagnosed in
the clinic or in the field [4–13], the diagnostician is not
always the person doing the corrective surgery, therapy,
or management. Cases are often transferred between or
among veterinarians and staff and, therefore, records
that contain terms that are out-dated and indefinite can
potentially result in misinterpretation of the original
diagnosis and, consequently, animals may receive
unintended treatment, which can be an economic loss
to the producer and, perhaps a liability to the
practitioner. Comprehensible and precise definitions
are necessary for differential and accurate diagnoses of
any disorder and for evaluating the benefit-to-risk ratio,
therapeutic regimes [14]. Correspondingly, scientists
involved in applied and basic research in cattle require a
lexicon of terms that is used uniformly among
individuals so that inference of the results between
and among studies can be correctly substantiated or
negated, and new hypotheses can be formulated without
unnecessary redundancy in experiments. Thus, defining
and adopting terminology used in bovine reproduction
that is clear, precise and understandable is expected to
make the exchange of clinical and research information
and outcomes more effective, safe and economical.
In 1976, a committee was convened and charged
with making recommendations for standardizing terms
used in bovine reproduction [15]. Although some of the
proposed terminology may be out-dated, the final report
recommended nomenclature that covered fetal mem-
branes, embryonic and perinatal periods, reproductive
dysgenesis and normal development of the bovine
conceptus. Over the last 20 years, numerous reports
[16–24] have become available that have defined
healthy and diseased conditions pertaining to the
principal organs (ovaries and uterus) of the bovine
reproductive tract during various reproductive states, as
well as rationale for correcting or managing the
disorders; however, the terminology is scattered
throughout the literature. There is no comprehensive
report that has focused on compiling the various terms
and definitions used to characterize ‘‘normal’’ and
‘‘abnormal’’ conditions associated with either the
ovaries or uterus into a single source.
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
The objective of this review is focused on the bovine
ovary and is expected to be one of a series of reviews to
summarize classical and current terminology that has
been used to define healthy and diseased conditions
associated with follicular and luteal gland development
in dairy and beef cattle, encompassing both Bos taurus
and Bos indicus cattle, and primarily within the context
of the estrous cycle.
2. Estrous cycle
Although terminology associated with the bovine
ovary will primarily be discussed in context with the
estrous cycle, most terms are also applicable to
pregnancy, post-partum and anestrus. Basic concepts
and terminology of the bovine estrous cycle will be
presented and summarized as previously reviewed [25]
to provide a framework for the discussion of the various
terms applied to ovarian dynamics during the cycle, as
well as during other reproductive states.
Terms used to describe the estrous cycle in cattle
have slightly different meanings and spellings. The
word ‘‘estrous’’ is an adjective used to describe the
estrous cycle or some aspect of it (e.g., estrous
behavior), whereas the word ‘‘estrus’’ is a noun
indicating the period of sexual receptivity. In regard
to the latter, the term ‘‘heat’’ is often used synony-
mously with estrus. In British and European literature,
‘‘oestrous’’ and ‘‘oestrus’’ are the preferred spellings,
based on the Greek origin of the word oestrus; a family
of biting insects (oestridae) that tormented cattle and
induced a state of frenzy. Estrual is also an adjective
Fig. 1. Terms and concepts related to stages within the follicular and luteal phases in association with respective changes in circulating
concentrations of estradiol (E2), progesterone (P4) and preovulatory (OV) gonadotropin surge during the bovine estrous cycle. Adapted from [25] and
reprinted with permission.
1345
used to identify a condition related to estrus (e.g.,
estrual fluid).
European and Zebu breeds of cattle are generally
considered seasonally polyestrus, which is defined as
the repeated distribution of estrous cycles throughout
the year. If the regularity of estrous cycles is interrupted
by pregnancy and lactation or, perhaps, by improper
nutrition, environmental stress or general and repro-
ductive diseases, there is a period of anestrus (without
cyclicity) or a condition of irregular estrous cycles. The
clinical importance for identifying the basis of anestrus
is discussed in detail in Section 4. It should be
recognized that polyestrus and anestrus can also be used
as adjectives (i.e., polyestrous and anestrous) preceding
a noun (e.g., anestrous cow).
Although puberty is generally defined as the life-
stage where the female has attained the capacity to
successfully produce offspring, the first ovulation in the
life of the animal marks the beginning of a rhythmic
pattern of reproductive cyclicity. The basis for the
estrous cycle is to provide females repeated opportu-
nities to become pregnant, as each estrous cycle usually
terminates with ovulation at the end of estrus. In
general, the length of the estrous cycle in beef and dairy
cattle ranges from 18 to 24 d [25], which is influenced,
in part, by the length of the luteal phase and number of
follicular waves per cycle as discussed in Section 3. The
length of the estrous cycle can be referenced from either
the detection of estrus to detection of the next estrus
(i.e., estrous cycle; Fig. 1) using visual observation or
marking devices, or from detection of ovulation to
detection of the next ovulation (i.e., interovulatory
1346 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
Fig. 2. Terms and concepts related to follicular wave emergence (WE)
and subsequent development of Wave 1 (anovulatory) and Wave 2
(ovulatory, OV) in association with respective FSH surges during the
bovine estrous cycle. Adapted from [1] and reprinted with permission.
interval; Fig. 2) using transrectal palpation or ultra-
sonography. In earlier studies, day of detection of estrus
was often used as a point of reference designated Day 1
and, in later studies, especially with the use of
ultrasonography [1], detection of ovulation has been
preferred as a less variable and more distinctive event to
serve as a point of reference, designated Day 0 as
indicated in Figs. 1 and 2. Through palpation or
ultrasonic imaging of the ovaries, ovulation is readily
detectable and is the recommended point of reference
that should be defined (i.e., Day 0 = ovulation) for
communicating the number of days post-ovulation in
non-pregnant as well as in pregnant animals [2]. In
situations where transrectal palpation experience and
ultrasound examinations are limited, the beginning of
observed estrus may still be used as reference; however,
it should be defined (e.g., Day 0 = first detection of
estrus), accordingly to avoid miscommunication of the
timing of therapies or treatments and misinterpretation
of study results.
When making comparisons between or among
studies that have used Day 0 or Day 1 relative to
estrus and Day 0 relative to ovulation as the reference
point, it may be necessary to adjust the data by 1 d so
that an approximation can be made to the day of
ovulation for a more accurate interpretation of the
results. In general, the time of ovulation is defined as the
disappearance of a dominant follicle that was present at
one examination and gone the next, as determined by
transrectal palpation or ultrasonography. Considering
that examinations are done daily or at approximately
24-h intervals, ovulation could occur soon after the last
examination, soon before the next examination or
anywhere in-between (i.e., within approximately 24 h
from the last examination). Furthermore, ovulation
typically occurs approximately 1 d after estrus
(approximately 29 h; [25]). Hence, by adjusting the
data relative to estrus by 1 d so that Day 1 or Day 2 post-
estrus is equivalent to ovulation on Day 0, a more
accurate assessment of the results between or among
studies using different times of reference can be made.
Physiologically and behaviorally, the estrous cycle
consists of the follicular phase with proestrus and estrus,
and the luteal phase with metestrus and diestrus, as
previously described [25]. Morphological, physiologi-
cal and endocrinological distinctions between the
various stages of the follicular and luteal phases of
the estrous cycle are summarized in the following
sections.
2.1. Follicular phase
The follicular phase is a relatively short period
(approximately 20%) of the estrous cycle, from the
beginning of regression of the corpus luteum to
ovulation, that involves proestrus and estrus (Fig. 1).
In general, the follicular phase is characterized by the
onset of estrous behavior and a shift from progesterone
dominance to estrogen dominance, primarily as a result
of structural and functional regression of a mature
corpus lutuem to a regressed corpus albicans (decreased
progesterone) and development of a preovulatory
follicle (increased estradiol) as depicted diagrammati-
cally (Figs. 1 and 2).
2.1.1. Proestrus
Proestrus (before estrus) is the initial portion of the
follicular phase that begins with structural and
functional regression of the corpus luteum and ends
at the onset of estrus (Fig. 1). As the luteal gland
regresses and circulating concentrations of progester-
one decrease, there is an increase in estradiol that plays
a role in the increase in uterine tone [26] and discharge
of clear and watery mucus from the vulva [27].
Follicular dynamics during proestrus involve continued
regression of the dominant follicle (Table 1) from the
previous anovulatory wave that overlaps with growth of
the dominant follicle of the ovulatory wave (Fig. 2).
2.1.2. Estrus
Estrus is the latter portion of the follicular phase that
ends with ovulation of the dominant follicle. Circulating
concentrations of estradiol reach peak concentrations
(Fig. 1) in association with the onset of sexual
receptivity, as indicated by estrous behavior (heat)
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357 1347

Table 1
Terms and concepts of ovarian follicles and corpora lutea associated with folliculogenesis and luteogenesis in cattle.
Cohort of follicles
Largest follicle
Second, third, fourth, etc.,
largest follicles
Dominant follicle
Subordinate follicles
Pre-ovulatory follicle
Corpus luteum
Corpus hemorrhagicum
Corpus luteum with a cavity
Corpus albicans
A cohort of follicles is a group of antral follicles that constitute a follicular wave from first detection
at 1–4 mm in diameter within a 2–3 d interval.
The largest follicle is the largest antral diameter follicle of the cohort of follicles of a wave before
deviation and is sometimes referred to as the future dominant follicle, due to its propensity to maintain
its hierarchal position within the cohort during the common growth phase and become the dominant follicle
after deviation.
The second, third, fourth, etc., largest follicles are the next largest follicles of the cohort of follicles of a
wave before deviation and are sometimes referred to as future subordinate follicles, due to their propensity
to maintain their hierarchal positions within the cohort during the common growth phase and become
respective subordinate follicles after deviation.
The dominant follicle is the largest antral follicle of the cohort of follicles of a wave after deviation that
typically reaches 10 mm in diameter and includes co-dominant or multiple dominant follicles that can
be ovulatory as well as anovulatory.
The subordinate follicles are the next largest antral follicles of a wave after deviation that are
typically <10 mm in diameter and are anovulatory.
The pre-ovulatory follicle is the dominant follicle after deviation that is selected for final maturation
and ovulation (sometimes referred to as the Graafian follicle).
The corpus luteum or luteal gland is formed at the site of the previous dominant follicle after ovulation,
through the structural and functional transformation of theca and granulosa cells to luteal cells.
Transient morphological characterization of a corpus luteum or luteal gland that is formed at the site of
the previous dominant follicle immediately after ovulation that develops an intra-luteal blood-filled cavity.
The term ‘‘corpus luteum with cavity’’ replaces the term ‘‘cystic corpus luteum’’ as a non-pathological
luteal gland in which a distinct fluid-filled cavity central to luteal tissue formation persists after the
initial formation of a corpus hemorrhagicum.
The corpus albicans is the remnant of the luteal gland following structural and functional regression of
luteal cells.

and discharge of cohesive strands of mucus from the
vulva. Final growth and maturation of the preovulatory
follicle (Fig. 2) occurs in association with a decrease in
estradiol concentrations and a preovulatory gonado-
tropin surge primarily involving LH and to a lesser
extent FSH, which precedes ovulation by approxi-
mately 29 h [25].
Detection of mounting behavior associated with
estrus (heat) is displayed by approximately 90% of cows
[28] and is critical for determining the onset of standing
estrus which is closely related to the timing of ovulation
(e.g., approximately 29 h post-estrus). Standing estrus
or standing heat with an arching of the back (standing to
be mounted) is a mating posture termed ‘‘lordosis’’,
which characterizes the most definitive sign of estrus.
Secondary signs of estrus may be indicative that
sexual receptivity is approaching; however, they may
occur before, during, or after estrus. They include,
trailing, sniffing the genitalia and attempts to mount
other animals, mucus discharge and swelling and
reddening of the vulva, and increased bellowing,
restlessness, chin resting, and lip curling. Occasionally
pregnant cows exhibit signs of estrus, usually during
middle to late gestation.
The terms ‘‘silent estrus’’ (silent heat) and ‘‘silent
ovulation’’ are used to characterize the condition of
ovulation without estrus [29]. Silent heat is a condition
that has been mostly associated with post-partum
ovulations in lactating cows. The incidences of silent
ovulation at the first, second, third and fourth ovulations
post-partum were reported to be 83, 46, 13, and 0%,
respectively [30]. Some clinicians [31] prefer to classify
silent ovulation as a type of anestrus termed ‘‘sub-
estrus’’ in which the incidence has been reported to
range from 34 to 50% [31,32]. However, since silent
ovulation is associated with ovulation and not
anovulation, it seems more appropriate to address
silent ovulation as a separate condition apart from
anestrus.
2.2. Ovulation
Ovulation is the terminal event of estrus character-
ized by the rupture of a dominant follicle, evacuation of
follicular fluid, granulosa cells and oocyte, and early
development or formation of a corpus luteum at the
ovulation site. Although cattle are considered a
monotocous species that typically ovulate a single
dominant follicle, there is some degree of multiple
spontaneous ovulations that appears to be influenced by
a variety of factors (e.g., breed, parity, nutrition, and
reproductive status) [33]. In one study involving 1917
1348 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
high-yielding dairy cows [34], the double ovulation rate
was 16%, with 52.7% unilateral (42.5% on the left
ovary and 57.5% on the right ovary) and 41.5% bilateral
ovulations. Furthermore, the triple ovulation rate was
5.8%. The incidence of synchronous (same day) versus
asynchronous (>1 d apart) ovulations was not reported.
2.3. Luteal phase
The luteal phase is the longer period of the estrous
cycle, comprising approximately 80% of the cycle from
ovulation to regression of the corpus luteum that
involves metestrus and diestrus (Fig. 1). In general, the
luteal phase is characterized by a shift from estrogen
dominance to progesterone dominance [35] following
ovulation (decreased estradiol), and structural and
functional maturation of a corpus lutuem (increased
progesterone). In association with the periodic emer-
gence of follicular waves and corresponding FSH
surges, there are also low-level fluctuations (i.e.,
transient increases and decreases) in systemic concen-
trations of estradiol associated with follicular wave
development throughout the luteal phase (Fig. 1).
2.3.1. Metestrus
Metestrus is the immediate time after estrus or the
initial portion of the luteal phase that begins at ovulation
and ends at diestrus (Fig. 1). In this regard, metestrus is
sometimes referred to as early diestrus, which extends
to approximately 5 d after ovulation. At the ovarian
level, metestrus involves luteinization of follicle cells of
the previous dominant follicle into a corpus luteum
(Table 1). Luteinization is a complex process that
involves remodeling of stromal and vascular tissue and
cellular and biochemical transformation of follicle
androgen- (theca) and estrogen (granulosa)-producing
cells into luteal progesterone-producing cells [36]. In
cattle, early formation of the luteal gland is often
associated with the formation of a transient intra-luteal
blood-filled cavity resembling a blood clot; this
structure is termed a ‘‘corpus hemorrhagicum’’
(Table 1) [25], which is comparable to the term used
in horses [41]. Thereafter, as the bovine luteal gland
completes its structural and functional transformation
towards the end of metestrus, it becomes morphologi-
cally homogeneous, appearing as a uniform echodense
structure upon detection with ultrasonography or
appears heterogeneous with an anechoic fluid-filled,
intra-luteal cavity [1]. Regarding the latter, intra-luteal
cavities in cattle can persist as fluid-filled cavities after
the initial stages of corpus hemorrhagicum formation
and range in size from small (e.g., <2 mm) to large
(e.g., >10 mm), such that the larger the cavity the
longer it is observed post-ovulation [37–39]. The
frequency of occurrence of one or the other morphol-
ogies (i.e., corpus luteum with or without a cavity) may
be different between heifers and cows. In cows [37], the
number of corpora lutea with a cavity versus without a
cavity was 37% versus 63%, whereas, in heifers [38],
the frequency was 79% versus 21%, respectively.
Despite the apparent difference between cows and
heifers in the occurrence of corpora lutea with and
without a cavity, progesterone concentrations, inter-
ovulatory intervals and pregnancy rates were not
affected by the different types of luteal gland
morphologies in cows and heifers [37,38].
Historically, a corpus luteum with an intra-luteal
cavity was referred to as a ‘‘cystic corpus luteum’’
which was considered a diseased or pathological
condition [40]. However, considering there is no
detectable difference in function (i.e., progesterone
output) or fertility (i.e., estrous cycle length and
pregnancy rate) between the two types of luteal gland
morphologies, it is generally accepted that a corpus
luteum with an intra-luteal cavity is not a pathological
condition. The term ‘‘cyst’’ or ‘‘cystic’’ can be used
anatomically in a pathological as well as a non-
pathological manner. To avoid confusion between the
terms ‘‘luteal cyst’’, which is considered a diseased,
morphological structure (see Section 5), and ‘‘cystic
corpus luteum’’, which is considered a healthy,
morphological structure, the contemporary term ‘‘corpus
luteum with a cavity’’ (Table 1) has been suggested to
replace the classical term ‘‘cystic corpus luteum’’ [37].
In association with an increase in the progesterone-
to-estrogen ratio during metestrus (Fig. 1), the
endometrial epithelium is denuded [42], particularly
in the intercaruncular areas [43], leading to the
appearance of bloody mucus at the lips of the vulva.
Metestrous bleeding is a process of diapedesis and is
characterized by a discharge of bloody mucus from the
vulva [43,44]. Metestrous bleeding is noticed 24–48 h
after estrus and is observed in 75–90% of heifers and
48–61% of cows [42], and its occurrence has no
correlation with conception. Metestrus also encom-
passes some aspects of development of the first
anovulatory wave (Wave 1) of the estrous cycle (Fig. 2).
2.3.2. Diestrus
Diestrus begins after metestrus and ends around the
beginning of regression of the corpus luteum, near the
time of proestrus (Fig. 1). The end of metestrus is
characterized by cessation of metestrous bleeding,
suppressed estrous behavior and a relatively mature
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
corpus luteum, with increasing systemic concentrations
of progesterone. Throughout diestrus, a mature corpus
luteum maintains maximal concentrations of progester-
one during emergence of Wave 2 (Fig. 2) and Wave 3 in
two- or three-wave estrous cycles.
In the absence of pregnancy, structural and func-
tional regression of the corpus luteum occurs sponta-
neously primarily in response to uterine PGF2a during
late diestrus [45]. Historically, the term ‘‘luteolysis’’ has
been most commonly used to describe regression or
lysis of the luteal gland. However, it appears to have a
very limited meaning that no longer describes properly
the complex sequence of synchronized molecular
events that are now known to be associated with the
demise of the corpus luteum. Contemporarily, therefore,
‘‘luteal regression’’ is the preferred terminology [46]
that may be preceded by the terms ‘‘functional’’ and/or
‘‘structural’’ to clarify the context in which luteal
regression is presented (i.e., endocrinologically or
morphologically).
Nonetheless, subsequent to diestrus, the corpus
luteum regresses structurally (e.g., decrease in dia-
meter) and functionally (e.g., decrease in progesterone)
in non-pregnant animals to form a corpus albicans
(Table 1) at the beginning of proestrus.
3. Follicular wave dynamics during the estrous
cycle
Follicular wave dynamics during the estrous cycle
has been well described in cattle and recently reviewed
[47–49]. The follicular wave pattern depicted in Fig. 2 is
that of a two-wave pattern; however, three-wave
patterns are also common in cattle (>95% of the
estrous cycles have two- or three-wave patterns), with
one- and four-wave patterns being rare [50,51]. In
general, development of a follicular wave involves the
organized and simultaneous growth of a cohort
(Table 1) of gonadotropin-dependent antral follicles
(i.e., wave emergence) within a 2–3 d interval. The
cohort of follicles associated with the wave initially
increase in size during a common growth phase and
subsequently differentiate into a single dominant
follicle (Table 1) that continues growth, whereas
multiple subordinate follicles (Table 1) cease growth
during a static phase (i.e., deviation; Fig. 2). By the time
the dominant anovulatory follicle of Wave 1 (and Wave
2 in three-wave estrous cycles) or the dominant
ovulatory follicle of Wave 2 or Wave 3 reaches its
static or preovulatory phase, corresponding subordinate
follicles have committed to regression. Subsequent to
deviation, morphological (i.e., size) and physiological/
1349
endocrinological (i.e., FSH-suppressing capacity) dom-
inance of the selected follicle is maintained for the
remainder of the growth phase and mid-way through the
static phase for anovulatory dominant follicles or until
ovulation. Although the physiological/endocrinological
dominance of the anovulatory follicle begins to wane
during the static phase, morphological dominance is
still apparent until regression to <10 mm (Fig. 2). In
this regard, more than one examination (e.g., transrectal
palpation or ultrasonography) is required to differenti-
ate between a growing and regressing follicle. As a
result of the loss of physiological/endocrinological
dominance at ovulation, an FSH-surge precedes
emergence of Wave 1 of the next cycle and, as a result
of a similar loss of dominance during the static phase, an
FSH-surge precedes emergence of Wave 2 in two-wave
estrous cycles (Fig. 2) and Wave 3 in three-wave cycles.
Studies using transrectal ultrasonography have
characterized the composition and nature of follicular
waves, and clarified the morphological differences and
timing of follicular events associated with multiple
waves in prepubertal and adult cattle during the estrous
cycle and pregnancy [1]. Breed, nutrition, parity and
lactation are some of the factors that influence the
number of follicular waves per estrous cycle, number or
cohort of follicles associated with a wave, and diameter
of the largest or future dominant follicle at deviation,
dominance and pre-ovulation.
The endocrinology of follicular wave dynamics has
been reviewed [49]; the most notable systemic
hormonal change is an FSH surge that stimulates the
emergence of each follicular wave during the estrous
cycle (Fig. 2). The initial increase in FSH has been
associated with follicles 1–3 mm in diameter [52], with
peak concentrations occurring around the time the
largest follicle of the wave reaches approximately 4 mm
[1]. Thereafter, FSH concentrations progressively
decrease, reaching basal concentrations during the late
growth/early static phase of the dominant follicle. The
next FSH surge and corresponding follicular wave soon
follow. Discovery of the FSH surge-follicular wave
interrelationship has led to subsequent discoveries on
the nature of follicle selection (e.g., deviation) which
have provided practical information for revising regi-
mens for hormonal synchronization of estrous cycles
used in breeding programs and timing of gonadotropin
treatments used in embryo transfer programs [49].
For hypothesis testing, major aspects of follicular
wave dynamics (i.e., emergence, deviation and dom-
inance) have involved the sequential tracking of
individual follicles to differentiate growing follicles
of one wave from regressing follicles of a previous wave
1350 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
[53]. For practical purposes, the major aspects of
follicular wave dynamics may be characterized by using
a nonidentity method to sequentially monitor and
determine the size of the three largest follicles within
each ovary [1]. Morphological characteristics of
follicular wave emergence, deviation and dominance
associated with follicular wave dynamics have been
recently reviewed [49] and are summarized in the next
sections.
3.1. Follicular wave emergence
Determining the day of wave emergence relative to
ovulation on Day 0 can be done prospectively or
retrospectively, according to when the largest follicle or
dominant follicle, respectively, of the wave first reaches
a prescribed diameter [49,54]. Based on when the
largest or dominant follicle of a wave first reached
4 mm, emergence of Wave 1 (anovulatory) and Wave
2 (ovulatory) in two-wave estrous cycles occurred on
approximately Days 0 and 10, respectively (Fig. 2),
whereas emergence of Wave 1 (anovulatory), Wave 2
(anovulatory) and Wave 3 (ovulatory) in three-wave
cycles occurred on approximately Days 0, 9 and 16,
respectively. With advanced ultrasound technology,
follicles 1–3 mm can be reliably and consistently
detected [49]. Hence, the day of wave emergence would
be expected to occur earlier (i.e., before ovulation) if
follicles of the wave are first detected at a smaller
diameter [52].
Wave emergence is associated with a hierarchal
arrangement of follicles, such that the largest follicle of
the cohort usually maintains its position throughout the
common growth phase and becomes the dominant
follicle a majority of the time [54]. In this regard, the
largest follicle is often referred to as the future dominant
follicle (Table 1) and the next largest follicles as future
subordinate follicles (Table 1) prior to deviation.
The interrelationship between the length of the luteal
phase and maximal diameter of dominant anovulatory
follicles influences the interwave interval (i.e., emer-
gence of one wave to emergence of the next wave;
Fig. 2) during the estrous cycle and, consequently, the
length of the cycle. While the estrous cycle or
interovulatory interval is significantly longer in cattle
with three- versus two-wave cycles (approximately 23 d
versus 20 d, respectively) as a result of a significantly
longer luteal phase (approximately 19 d versus 16 d,
respectively), the interwave intervals are shorter in
cattle with three-wave cycles, in association with
smaller dominant anovulatory and ovulatory follicles
compared to cattle with two-wave cycles [1,50].
Notably, spontaneous or hormonally induced prolonga-
tion of the luteal phase has been associated with Type IV
anestrus and is discussed in Section 4.
3.2. Follicle selection and deviation
In general, follicle selection is a process where, in
monotocous species, typically one follicle is selected
from a cohort of growing follicles and becomes
dominant, with continued growth to preovulatory size
before regressing or ovulating (Fig. 2); the remaining
follicles of the wave (i.e., subordinates) regress. The
follicle selection process is often thought to be a single
event associated with the differential change in growth
rates between the largest and next largest follicles of a
wave (i.e., follicle deviation). However, the selection
process likely involves several morphological and
physiological/endocrinological events, perhaps, begin-
ning with the settlement of primordial germ cells within
the early ovary of the embryo/fetus. Other events under
the umbrella of the follicle selection process may include
the timing of recruitment of primordial follicles from the
resting pool, hierarchal arrangement in growth of
preantral follicles, stage of development, and size of
antral follicles at the time of wave emergence, and
commitment of subordinate follicles to regression
following the differential change in development
(morphological, physiological and endocrinological) of
the largest and next largest follicles of a wave [55–65].
To distinguish the abrupt separation in growth
between the dominant and largest subordinate follicles
in individual animals [56] during the selection process
from the gradual separation in the mean diameter
profiles averaged among animals [50], the term
‘‘follicle deviation’’ was coined to replace ‘‘follicle
divergence’’, respectively. The timing of follicle
deviation within an animal can be determined sub-
jectively by visual assessment of the diameter growth
profiles between the two largest follicles [62]. The point
of departure, or the time of the differential or distinctive
change in the diameter profiles, represents follicle
deviation. The visual approach was compared to a more
objective approach using a segmented regression model
in Bos taurus [53] and Bos indicus [64] cattle. There was
no difference between the two methods in either
species; hence, the visual approach was considered
reliable and more practical. Thus, follicle deviation
occurs approximately 2.5 d after wave emergence (i.e.,
largest follicle 4 mm) and represents a major and
readily observable morphological event involved in the
follicle selection process associated with ovulatory as
well as anovulatory follicular waves in cattle.
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
The morphological and physiological/endocrinologi-
cal basis for follicle deviation has been reviewed in cattle
[65–70]. In general, after wave emergence and peak
concentrations of FSH, the common growth phase of
wave development is associated with the declining
portion of the FSH surge. During this time, there are
differential changes between the future dominant and
subordinate follicles regarding local (e.g., IGF and IGF
binding proteins) and systemic hormonal mechanisms
that, in part, regulate the decrease in FSH concentrations
[59,62]. By the end of the common growth phase, there is
a differential change in growth rates between the future
dominant and subordinate follicles (i.e., follicle devia-
tion). The time of deviation in Bos taurus non-lactating
dairy and beef cattle according to diameter of the largest
follicle has been reported to be approximately 8 mm [33]
and, in lactating Holstein cows, approximately 9 mm
[33]. In Bos indicus cattle, deviation occurred when the
largest follicle reached approximately 6 mm [64]. The
differences among cattle with respect to the timing of
follicle deviation according to follicle diameter provides
practical information for scheduling gonadotropin treat-
ments to avoid the potential confounding, suppressing
effect of follicle dominance on development of multiple
dominant follicles (i.e., superstimulation) and ovulations
(superovulation) [49].
3.3. Follicle dominance
Subsequent to deviation, morphological dominance
can be observed by the size of the dominant follicle,
using transrectal palpation or B-mode ultrasonic
imaging [1], physiologically by blood-flow using
color-Doppler ultrasonic imaging [3], and endocrino-
logically by follicular-fluid hormone concentrations
using immunoassays [59]. Physiological/endocrinolo-
gical dominance has been substantiated by the negative
influence the dominant follicle has on subordinate
follicles of the extant wave and emergence of
subsequent waves using follicle ablation and gonado-
tropin and follicular fluid treatments [47,49,56,62].
In general, a dominant follicle has been defined by size
when it reaches 10 mm [1,62]. Dominance could also
be defined according to its size at the time of deviation or
its ovulatory capacity to respond to an ovulatory
stimulus, as reported in Bos taurus [71] and Bos indicus
[72] cattle. Notably, multiple dominant follicles may be
associated with a wave, whether they are spontaneous or
gonadotropin-treatment induced. The maximum dia-
meter of the dominant follicle of anovulatory and
ovulatory waves is approximately 16 mm [56] in Bos
taurus and 11 mm in Bos indicus [73]. The nature of
1351
dominance represents both physiological (e.g., vascu-
larity, granulosa cell LH receptors) and endocrinological
(e.g., follicular fluid estradiol and inhibin) aspects of the
dominant follicle during its growing phase, that indirectly
influence the regression of subordinate follicles and
suppression of subsequent wave emergence through
suppression of FSH (Fig. 2). The sequence of events
associated with periodic surges of FSH and correspond-
ing follicular waves is repeated throughout the estrous
cycle and even into pregnancy, as indicated by the
continuous emergence of follicular waves during
gestation [1]. The regularity of these events, however,
may be interrupted by disease conditions associated with
the ovaries, which are addressed in the next sections.
4. Classification of anestrus
Anestrus is a broad term that indicates the lack of
expression of estrus (or absence of estrous signs), despite
a diligent estrus detection approach. Although absence of
behavioral and physiological characteristics of estrus can
fall under anestrus, a true anestrous condition is primarily
characterized by anovulation. Historically, anestrus was
broadly classified into physiological and pathological
(clinical) types, with the following representing the
pathological type: (1) inactive ovaries (i.e., minimal
follicle development, anovulation and no corpus luteum
development); (2) silent ovulation (i.e., ovulation without
behavioral estrus); (3) ovarian hypofunction (i.e.,
persistent dominant follicle); (4) cystic ovarian degen-
eration (i.e., follicular or luteinized-follicular cyst); and
(5) persistent corpus luteum (i.e., lack of luteal
regression) [74]. Contemporarily, anestrus has been
classified in a novel manner according to ovarian
follicular and luteal dynamics [23]. As discussed in
the previous section, follicular wave dynamics involves
three main morphological events: (1) emergence, (2)
deviation, and (3) dominance ending in anovulation or
ovulation. Classification of anestrus or anovulation based
on follicle characteristics at emergence, deviation and
dominance provides for a rational diagnosis and
treatment of the underlying diseased condition. It should
be pointed out that silent ovulation (lack of overt signs of
estrus) and unobserved estrus (poor estrus detection
approach) can apparently increase the incidence of
anestrus in a herd; however, they are not included in this
classification since the former is behavioral and the latter
is a management issue. Only true or organic forms of
anestrus [75] are classified in this review.
For discussion purposes, the etiologies of the
following four types of anestrus are depicted dia-
grammatically (Fig. 3).
1352 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
Fig. 3. Schematic representation of types of anestrous conditions based on the morphology and physiology of ovarian follicles in cattle. Adapted
from [83] and reprinted with permission.
Type I: Follicle growth proceeds to detection of
antral follicles at wave emergence and to pre-deviation
size (e.g., follicles <10 mm) but not to dominance (i.e.,
follicle 10 mm). Diagnosis is based on multiple
transrectal ultrasound examinations indicating follicles
<10 mm in diameter and no detectable corpus luteum.
Hence, Type I anestrus corresponds to the earlier
classification of anestrus due to relatively inactive
ovaries according to reduced follicle growth [74].
Type II: Follicle deviation and dominance is attained
but the dominant follicle of the ovulatory wave fails to
ovulate and, instead, regresses. Diagnosis is based on
multiple transrectal ultrasound examinations at 7-d
intervals, indicating no detectable corpus luteum or
follicular cyst [76,77]. Similarly, Type II anestrus also
corresponds to the earlier classification of anestrus due
to relatively inactive ovaries and failure of ovulation
[74,77].
Type III: Follicle dominance is achieved, but the
dominant follicle of the anovulatory or ovulatory wave
fails to regress or ovulate at the expected time and
remains at preovulatory size or increases in size to form a
persistent dominant follicle [78,79]. Persistent dominant
follicles may develop into a follicular cyst or become
luteinized to form luteinized-follicular cyst (i.e., cystic
follicular degeneration; Section 5). These anovulatory
structures may persist for an extended interval before
eventually regressing. Depending on the functional status
of these structures during their development, they may be
associated with the suppressed emergence of another
follicular wave. Diagnosis is based on the history and
morphological characteristics of follicular and luteal
dynamics. Hence, Type III anestrus corresponds to the
earlier classifications of anestrus due to ovarian
hypofunction and cystic ovarian degeneration, COD
[74]. The pathological nature of COD and discussed in
more detail in Section 5.
Type IV: Although estrus is exhibited and ovulation
and formation of a corpus luteum occurs as usual, the
luteal gland persists beyond the expected time of
regression, resulting in anestrus or an extended inter-
estrus or interovulatory interval. A contributing factor
may be the absence of an estrogenic dominant follicle at
the expected time of regression of the corpus luteum [23].
Elevated systemic estradiol concentrations associated
with development of the dominant follicle reportedly
lead to an increase in uterine estradiol receptors and
up-regulation of uterine oxytocin receptors and,
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
consequently, pulsatile release of PGF2a [45]. In regard to
the latter, uterine infection [80] and related pyometra can
prolong the life of a corpus luteum as a result of reduced
uterine PGF2a production [81]. Parity, dystocia, puerp-
eral disturbances, premature lactation, heat stress, and
early resumption of ovarian cyclicity after calving are
other factors that have been suggested to increase the
persistence of a corpus luteum and prolongation of the
luteal phase [82]. Diagnosis is based on the history and
morphological characteristics of the ovaries and uterus.
Type IV anestrus (i.e., lack of luteal regression)
corresponds to the earlier description of anestrus due
to a persistent corpus luteum [74].
Treatment options for these four types of anestrus
conditions are beyond the scope of this discussion but
have recently been reviewed [83].
5. Ovarian diseases
The ovaries play a key role in reproduction as the
source of oocytes and steroid hormones. Estrogens and
progestins are necessary for behavioral and physiolo-
gical aspects associated with follicular development,
ovulation, luteal gland formation, pregnancy main-
tenance, parturition and lactation. Impairment of the
hormonal control of any of these reproductive events
can lead to acute or chronic sterility or infertility issues.
Although gross anatomical images of some of the
ovarian conditions discussed in the next sections have
been complied [84], a compilation of ultrasonographic
images of diseased or pathological conditions of bovine
ovaries is lacking.
5.1. Follicular and luteinized-follicular cysts
Follicular and luteinized-follicular cysts are collec-
tively referred to as abnormal structures associated with
cystic ovarian degeneration or disease. As mentioned
earlier (Type III anestrus), a follicular cyst arises from a
persistent dominant follicle. Although the etiology is not
thoroughly known, if left undiagnosed, the follicle cells
(granulosa and theca) of the cyst begin to luteinize, such
that they take on different structural and functional
characteristics to become a luteinized-follicular cyst
previously known as a luteal cyst [85]. A follicular cyst is
defined as an anovulatory follicle-like structure [86] of
the ovary which is fluid-filled, usually >24 mm in
diameter, which persists for more than 7–10 d in the
absence of a corpus luteum [87]. In regard to the latter, the
presence or absence of a corpus luteum as a component of
the definition of a luteinized-follicular cyst is debatable
[88]. In addition, the size of a luteinized-follicular cyst
1353
may be <20 mm in experimentally induced cysts,
particularly, if there is more than one [24].
Based on transrectal palpation, it has been estab-
lished that follicular cysts are typically larger (>2 cm)
than a preovulatory-sized follicle and have a feeling of
being thin walled, fluid-filled and easily ruptured during
manipulation, whereas luteinized-follicular cysts have a
firmer feel and are less fragile, due in part, to a wall of
apparent luteal tissue. According to ultrasonographic
morphology, both follicular and luteinized-follicular
cysts appear as anechoic structures; however, the latter
appears with a thicker wall of apparent luteal tissue [87]
that can vary from an irregular pattern to a fairly well-
defined border [90,91]. Morphologically, luteinized-
follicular cysts may resemble corpora lutea with a
cavity and, endocrinologically, circulating concentra-
tions of progesterone are similar to concentrations
during the expected luteal phase of the estrous cycle
[92,93]. Although ultrasonography has provided a
better understanding of the characteristics associated
with gross development, diagnosis and differentiation
between follicular and luteinized-follicular cysts, the
etiology [85] and optimal regimens for treatment of the
condition [87,89] are not thoroughly known.
5.2. Parovarian cysts
Parovarian cysts are cystic structures near the ovaries
in the broad ligament and close to the uterine tubes.
They can be detected using transrectal palpation or
ultrasonography as fluid-filled, anechoic structures and
are usually round or oval in shape, occur as a single
cystic structure, and range from 1–5 cm in diameter.
The relatively larger paraovarian cysts are referred to as
hydatid of Morgagni, similar to those observed in ewes
and sows [94,95]. Paraovarian cysts are vestiges of the
mesonephric (Wolffian) or paramesonephric (Muller-
ian) duct systems and are of two types. Cysts derived
from the cranial mesonephric tubules are referred to as
epoophoron, whereas those derived from the caudal
mesonephric tubules are referred to as paroophoron.
Paraovarian cysts are considered morphologically and
physiologically benign; they have no apparent effect on
the estrous cycle and fertility [96].
5.3. Fibrin tags
Small tags of fibrin, particularly in heifers, from
bleeding after ovulation are frequently attached to the
ovary at the site of a previous ovulation or on the medial
attachment of the ovary to the uteroovarian ligament
and are commonly referred to as ovulation tags. There is
1354 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
a paucity of information about these tags, but it appears
that they do not interfere with subsequent ovulations or
conceptions [96].
5.4. Granulosa-theca cell tumor
While ‘‘granulosa cell tumor’’ has been the classic
terminology, ‘‘granulosa-theca cell tumor’’ is the pre-
ferred terminology since it appears both types of follicle
cells are involved in the disease. Even though granulosa-
theca cell tumors are the most common ovarian tumors in
cattle, they are rare (<0.5%). This type of tumor arises
from the sex cord stromal tissue within the ovary and may
be relatively small, solid, and yellow to white or large,
filled with cysts of varying sizes and weigh 11.9–12.3 kg
[96]. They are usually benign (no metastasis) but can be
malignant and are often hormonally active. If undiag-
nosed or diagnosed and left intact, clinical signs progress
through various stages beginning with nymphomania and
ending with virilism. In some instances, mammary
development is observed [97].
Diagnosis is based on clinical signs and transrectal
palpation and ultrasound examinations [86]. Clinically,
granulosa-theca cell tumors may be characterized by
abnormal estrous cycles and follicular and luteal
inactivity on the contralateral ovary. In the mare,
granulosa-theca cell tumors are also differentiated by
assay of circulating concentrations of androgens and
inhibin [98], which apparently have not been fully
characterized as diagnostic tools for granulosa-theca
cell tumors in cattle.
5.5. Rete ovarii (ovarian cyst)
In the hilus region of a mature ovary, the rete ovarii is
found as a network arrangement of medullary tubules or
cords near the mesoovarium. The cords are lined with
cuboidal and columnar epithelium that differentiate to
form granulosa-like cells possessing secretory activity
[99]. Cysts associated with rete ovarii are not commonly
linked to severe pathological lesions [100,101]; how-
ever, if the rete ovarii has a space occupying lesion,
ovarian function may be jeopardized. Transrectal
ultrasonic imaging of the reproductive tract has
provided a tool for diagnosing and differentiating rete
ovarii from other structures associated with the ovary.
5.6. Ovarian bursa adhesions
A perivoarian adhesion usually results from ovarian
trauma or peritonitis that may lead to blockage of ovum
transport from the ovaries and embryo migration through
the uterine tubes. Historically, adhesions have been
associated with physical rupture of a large cyst or
expression of a corpus luteum. Fortunately, these
antiquated procedures are no longer commonly practiced.
Contemporarily, uterine irrigation with irritating fluids in
large volumes for therapeutic and non-therapeutic reason
may leak into the bursa area and provoke an inflammatory
response. Notably, intensive and repeated handling of the
reproductive organs involved in extensive reproductive
programs may induce microscopic lesions on the
peritoneal surface. In one instance involving 50% of
the cows, lesions were present in the uterine tubes and
adjacent tissues. These strands were made up of numerous
filaments, probably of collagen, and most likely
correspond to the organized peritoneal fibrinous exudate
that forms during peritonitis [102].
Adhesions between the mesosalpinx and mesoovar-
ium often include the fimbria and ovary and may be
mild, consisting of a few fibrous strands, moderate with
more than a few strands but not enough to interfere with
ovulation, or extensive to the point of concealing the
ovaries which can interfere with ovulation. In extreme
cases, adhesions may extend to the opening of the
ovarian bursa, resulting in a very narrow opening that
may affect fertility [103]. This condition is referred to as
perisalpingitis and is rare [104].
Diagnosis is based on reproductive history and
previous therapy for reproductive disorders combined
with detailed and methodical transrectal and ultrasono-
graphic examination of the reproductive tract.
5.7. Ovarian hypoplasia
The hypoplastic ovary undergoes incomplete devel-
opment (ovarian dysgenesis), such that the ovary lacks its
usual full complement of primordial follicles. Hypopla-
sia can be unilateral or bilateral and can be partial or
complete. Historically, this condition has been observed
and documented in certain breeds of dairy and beef cattle.
In heifers, hypoplastic ovaries may be small such
that they are difficult to locate by transrectal palpation.
The ovaries may feel like thin, narrow, firm cord-like
structures. Lack of estrus associated with this condition
has to be differentiated from anestrous conditions
associated with other ovarian disorders previously
discussed. Diagnosis is primarily based on transrectal
examination.
6. Summary
As one of a potential series of reviews, this review has
focused on compiling many of the classical and current
 A.T. Peter et al. / Theriogenology 71 (2009) 1343–1357
terms used to define healthy and diseased conditions of
the bovine ovary primarily during the estrous cycle.
Although some classical terms associated with follicular
and luteal gland development are still relevant today,
other terms are antiquated (e.g., cystic corpus luteum,
cystic ovarian degeneration, luteolysis, granulosa cell
tumor) and are advocated herein to be replaced with more
cotemporary terms (e.g., corpus luteum with a cavity,
follicular and luteinized-follicular cysts, structural and
functional luteal regression, granulosa-theca cell tumor).
By compiling many classical and contemporary terms
associated with ovarian dynamics in cattle into a single
source that is clear, precise and understandable, it is
expected that this review will make the exchange of
clinical and research information and outcomes more
effective, safe and economical.
Acknowledgements
In thanks and in memory of Howard Levine whose
dedication and knowledge of bovine reproduction
initiated the processes leading to this review. The
authors also thank the following members of the
American College of Theriogenologists for their
valuable assistance in reviewing and contributing their
knowledge in bovine reproduction to this effort: Gregg
Adams, Louis Archbald, Lionel Dawson, Donald
Schlafer and William Thatcher.
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