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Topic:Photoperiod

Photoperiod refers to the duration of daylight and darkness that an organism is exposed to within a 24-hour period. In horses, photoperiod significantly influences physiological and behavioral processes, including reproductive cycles, coat growth, and metabolic functions. Horses are long-day breeders, meaning their reproductive activity is stimulated by increasing daylight, which affects hormone production and fertility. Additionally, changes in photoperiod can impact melatonin secretion, which in turn influences seasonal hair coat changes and overall energy metabolism. This page compiles peer-reviewed research studies and scholarly articles that explore the effects of photoperiod on equine physiology, behavior, and management practices.
Acute injection and chronic perfusion of kisspeptin elicit gonadotropins release but fail to trigger ovulation in the mare.
Biology of reproduction    February 20, 2014   Volume 90, Issue 2 36 doi: 10.1095/biolreprod.113.114157
Decourt C, Caraty A, Briant C, Guillaume D, Lomet D, Chesneau D, Lardic L, Duchamp G, Reigner F, Monget P, Dufourny L, Beltramo M, Dardente H.Kisspeptin has emerged as the most potent gonadotropin-releasing hormone (GnRH) secretagogue and appears to represent the penultimate step in the central control of reproduction. In the sheep, we showed that kisspeptin could be used to manipulate gonadotropin secretion and control ovulation. Prompted by these results, we decided to investigate whether kisspeptin could be used as an ovulation-inducing agent in another photoperiodic domestic mammal, the horse. Equine kisspeptin-10 (eKp10) was administered intravenously as bolus injections or short- to long-term perfusions to Welsh pony mares, ei...
Blue light from individual light masks directed at a single eye advances the breeding season in mares.
Equine veterinary journal    December 5, 2013   Volume 46, Issue 5 601-605 doi: 10.1111/evj.12153
Murphy BA, Walsh CM, Woodward EM, Prendergast RL, Ryle JP, Fallon LH, Troedsson MH.Artificial lighting is commonly used to advance the breeding season in horses. Light masks have been developed that direct light at a single eye to inhibit the production of melatonin, the decoder of photoperiod for seasonally breeding animals. Objective: To investigate whether low-intensity blue light from light masks was effective at advancing the breeding season in mares. Methods: Controlled experiment. Methods: Data on reproductive activity was collected from 3 groups of mares maintained on Kentucky horse farms under various lighting conditions between 20 November 2011 and 10 February 2012...
Contraception can lead to trophic asynchrony between birth pulse and resources.
PloS one    January 28, 2013   Volume 8, Issue 1 e54972 doi: 10.1371/journal.pone.0054972
Ransom JI, Hobbs NT, Bruemmer J.Abiotic inputs such as photoperiod and temperature can regulate reproductive cyclicity in many species. When humans perturb this process by intervening in reproductive cycles, the ecological consequences may be profound. Trophic mismatches between birth pulse and resources in wildlife species may cascade toward decreased survival and threaten the viability of small populations. We followed feral horses (Equus caballus) in three populations for a longitudinal study of the transient immunocontraceptive porcine zona pellucida (PZP), and found that repeated vaccinations extended the duration of in...
Investigation of a non-invasive method of assessing the equine circadian clock using hair follicle cells.
Journal of circadian rhythms    October 5, 2012   Volume 10, Issue 1 7 doi: 10.1186/1740-3391-10-7
Watts LM, Browne JA, Murphy BA.A comprehensive understanding of the equine circadian clock involves the evaluation of circadian clock gene expression. A non-invasive and effective method for detecting equine clock gene expression has yet to be established. Currently, research surrounding this area has relied on collecting tissue biopsies or blood samples that can often be costly, time consuming and uncomfortable for the animal. Methods: Five mares were individually stabled under a light-dark (LD) cycle that mimicked the external environmental photoperiod during a time of year corresponding with the vernal equinox. Hair foll...
Circulating pituitary and gonadal hormones in spring-born Thoroughbred fillies and colts from birth to puberty.
The Journal of reproduction and development    May 19, 2012   Volume 58, Issue 5 522-530 doi: 10.1262/jrd.2011-025
Dhakal P, Hirama A, Nambo Y, Harada T, Sato F, Nagaoka K, Watanabe G, Taya K.The present study was conducted to elucidate the profile of circulating gonadotropins and gonadal hormones from birth to puberty and relationship between gonadal seasonality and hormonal secretion in both sexes of Thoroughbred horses. Spring-born colts (n=6) and fillies (n=9) were blood sampled weekly from jugular vein from birth to 60 weeks of age. Circulating FSH, LH, prolactin, testosterone, progesterone, estradiol-17β, and immunoreactive (ir)-inhibin were measured by radioimmunoassay. In both sexes, the steroid hormones levels were remarkably high at birth, rapidly dropped within a week a...
Reproductive seasonality in the mare: neuroendocrine basis and pharmacologic control.
Domestic animal endocrinology    May 1, 2012   Volume 43, Issue 2 103-115 doi: 10.1016/j.domaniend.2012.04.001
Williams GL, Thorson JF, Prezotto LD, Velez IC, Cardoso RC, Amstalden M.Reproductive seasonality in the mare is characterized by a marked decline in adenohypophyseal synthesis and secretion of LH beginning near the autumnal equinox. Thus, ovarian cycles have ceased in most mares by the time of the winter solstice. Endogenous reproductive rhythms in seasonal species are entrained or synchronized as a result of periodic environmental cues. In the horse, this cue is primarily day length. Hence, supplemental lighting schemes have been used managerially for decades to modify the annual timing of reproduction in the mare. Although a full characterization of the cellular...
Effects of nutritional cues on the duration of the winter anovulatory phase and on associated hormone levels in adult female Welsh pony horses (Equus caballus).
Reproductive biology and endocrinology : RB&E    September 29, 2011   Volume 9 130 doi: 10.1186/1477-7827-9-130
Salazar-Ortiz J, Camous S, Briant C, Lardic L, Chesneau D, Guillaume D.Mares have an annual reproductive rhythm, with a phase of inactivity in midwinter. The aim of this study was to determine the impact of food restriction on physiological and metabolic hallmarks of this rhythm. Methods: Over three successive years, 3 groups of 10 mares were kept under natural photoperiod. A 'well-fed' group was fed to maintain the mares in good body condition; a 'restricted' group received a diet calculated to keep the mares thin and a 'variable' group was fed during some periods like the 'restricted' group and during some other periods like the 'well-fed' group, with the aim o...
Experimental jetlag disrupts circadian clock genes but improves performance in racehorses after light-dependent rapid resetting of neuroendocrine systems and the rest-activity cycle.
Journal of neuroendocrinology    September 17, 2011   Volume 23, Issue 12 1263-1272 doi: 10.1111/j.1365-2826.2011.02222.x
Tortonese DJ, Preedy DF, Hesketh SA, Webb HN, Wilkinson ES, Allen WR, Fuller CJ, Townsend J, Short RV.Abrupt alterations in the 24-h light : dark cycle, such as those resulting from transmeridian air travel, disrupt circadian biological rhythms in humans with detrimental consequences on cognitive and physical performance. In the present study, a jetlag-simulated phase shift in photoperiod temporally impaired circadian peaks of peripheral clock gene expression in racehorses but acutely enhanced athletic performance without causing stress. Indices of aerobic and anaerobic capacities were significantly increased by a phase-advance, enabling prolonged physical activity before fatigue occurred. Thi...
Absence of a serum melatonin rhythm under acutely extended darkness in the horse.
Journal of circadian rhythms    May 10, 2011   Volume 9 3 doi: 10.1186/1740-3391-9-3
Murphy BA, Martin AM, Furney P, Elliott JA.In contrast to studies showing gradual adaptation of melatonin (MT) rhythms to an advanced photoperiod in humans and rodents, we previously demonstrated that equine MT rhythms complete a 6-h light/dark (LD) phase advance on the first post-shift day. This suggested the possibility that melatonin secretion in the horse may be more strongly light-driven as opposed to endogenously rhythmic and light entrained. The present study investigates whether equine melatonin is endogenously rhythmic in extended darkness (DD). Methods: Six healthy, young mares were maintained in a lightproof barn under an LD...
Reproductive cycles of horses.
Animal reproduction science    February 18, 2011   Volume 124, Issue 3-4 220-228 doi: 10.1016/j.anireprosci.2011.02.005
Aurich C.Horses are long-day breeders. During the breeding season, cycle length is about 22 days with 5-7 days of oestrus. Gonadotroph cells are localized in the pars distalis as well the pars tuberalis of the pituitary and heterogeneity in the pattern of LH and FSH storage within the gonadotroph population is considered the basis for the differential regulation of gonadotrophin secretion throughout the reproductive cycle. No short and distinct periovulatory LH peak exists in the mare. The equine ovary has an extreme large size and weight. One to two major follicular waves develop per cycle. The preovu...
Clinical use of dopamine antagonist sulpiride to advance first ovulation in transitional mares.
Theriogenology    September 26, 2010   Volume 75, Issue 1 138-143 doi: 10.1016/j.theriogenology.2010.07.019
Panzani D, Zicchino I, Taras A, Marmorini P, Crisci A, Rota A, Camillo F.Artificial photoperiod treatment is currently the best method to hasten the first ovulation of the breeding season in winter anoestrus mares. However, this is not easy to apply in large herds of mares and, to be effective, has to be planned in the northern hemisphere in December at the latest. Pharmacological treatments have been proposed as alternatives: GnRH agonists, progesterone or its synthetic agonist Altrenogest, and dopamino-antagonists, as pherphenazine, domperidone or sulpiride. Dopamino-antagonists protocols, beginning at a given day of the year, gave controversial results in terms ...
The use of adrenocorticotrophic hormone as a potential biomarker of pituitary pars intermedia dysfunction in horses.
Veterinary journal (London, England : 1997)    June 12, 2010   Volume 185, Issue 1 58-61 doi: 10.1016/j.tvjl.2010.04.014
Lee ZY, Zylstra R, Haritou SJ.Elevated concentrations of plasma adrenocorticotrophic hormone (ACTH) in horses with pituitary disturbances are often associated with dysfunction of the pituitary's pars intermedia. The majority of such animals exhibit an increased susceptibility to laminitis, particularly during the autumn. The 24h plasma ACTH profiles of horses with pituitary pars intermedia dysfunction (PPID), and those of matched controls, were determined in March, June, September and December. Differences in ACTH concentrations between the groups were significant (P<0.0001), regardless of photoperiod, and persisted thr...
Role of prolactin in the gonadotroph responsiveness to gonadotrophin-releasing hormone during the equine annual reproductive cycle.
Journal of neuroendocrinology    March 2, 2010   Volume 22, Issue 6 509-517 doi: 10.1111/j.1365-2826.2010.01986.x
Hodson DJ, Townsend J, Gregory SJ, Walters C, Tortonese DJ.A combined suppressive effect of prolactin (PRL) and dopamine on the secretion of luteinising hormone (LH) at the level of the pituitary gland has been identified in sheep, a short-day breeder. However, little is known about the role of PRL in the intra-pituitary regulation of the gonadotrophic axis in long-day breeders. In the present study, we investigated the effects of PRL on LH and follicle-stimulating hormone (FSH) secretion during the equine annual reproductive cycle. Horse pituitaries were obtained during the breeding season (BS) and nonbreeding season (NBS). Cells were dispersed, plat...
Seasonal functional relevance of sperm characteristics in equine spermatozoa.
Theriogenology    January 18, 2010   Volume 73, Issue 7 950-958 doi: 10.1016/j.theriogenology.2009.11.023
Gamboa S, Rodrigues AS, Henriques L, Batista C, Ramalho-Santos J.A group of stallions with different reproductive indexes were used to study seasonal variations in sperm quality (Equus caballus). Semen samples were collected from late September to July and analyzed according to four seasonal periods: late September-December, January-March, late March-May, and June-July. Parameters monitored included sperm concentration, sperm motility, sperm morphology, sperm viability, acrosomal status, plasma membrane stability, and sperm mitochondrial membrane potential. Overall, seminal parameters monitored are affected mostly by time period, followed by animal and last...
Evaluation of plasma ACTH, alpha-melanocyte-stimulating hormone, and insulin concentrations during various photoperiods in clinically normal horses and ponies and those with pituitary pars intermedia dysfunction.
Journal of the American Veterinary Medical Association    September 16, 2009   Volume 235, Issue 6 715-722 doi: 10.2460/javma.235.6.715
Beech J, Boston RC, McFarlane D, Lindborg S.To measure plasma ACTH, alpha-melanocyte-stimulating hormone (aloha-MSH), and insulin concentrations during various photoperiods between February and October in horses and ponies with and without pituitary pars intermedia dysfunction (PPID). Methods: Cohort study. Methods: 13 clinically normal (control) ponies, 14 clinically normal (control) horses, 7 ponies with PPID, and 8 horses with PPID. Methods: Blood samples were collected from February through October during 8 photoperiods: 1, February 13 through March 2; 2, April 4 through 6; 3, June 19 through 22; 4, August 6 through 7; 5, August 14 ...
Gonadotrophin subunit and GnRH receptor gene expression in the pars distalis of the equine pituitary.
General and comparative endocrinology    December 10, 2008   Volume 160, Issue 3 236-242 doi: 10.1016/j.ygcen.2008.11.029
Townsend J, Westcott K, Tortonese DJ.In the horse, pronounced changes in fertility occur annually in response to photoperiod. However, the mechanisms regulating gonadotrophin synthesis and release in this species remain unclear. Here, we investigated the expression of gonadotrophin subunits and GnRH receptor (GnRH-R) mRNA in the pituitary glands of Thoroughbred horses during the breeding (BS) and non-breeding (NBS) season. Seasonal effects on the prevalence of gonadotrophs in the pars distalis were also examined. GnRH-R and common alpha-, LHbeta- and FSHbeta-subunit mRNA contents were determined by Northern analysis and the preva...
Seasonal variations in daily rhythms of activity in athletic horses.
Animal : an international journal of animal bioscience    July 1, 2008   Volume 2, Issue 7 1055-1060 doi: 10.1017/S1751731108002267
Bertolucci C, Giannetto C, Fazio F, Piccione G.Circadian rhythms reflect extensive programming of biological activity that meets and exploits the challenges and opportunities offered by the periodic nature of the environment. In the present investigation, we recorded the total activity of athletic horses kept at four different times of the year (vernal equinox, summer solstice, autumn equinox and winter solstice), to evaluate the presence of seasonal variations of daily activity rhythms. Athletic Thoroughbred horses were kept in individual boxes with paddock. Digitally integrated measure of total activity of each mare was continuously reco...
Seasonal changes in circadian peripheral plasma concentrations of melatonin, serotonin, dopamine and cortisol in aged horses with Cushing’s disease under natural photoperiod.
Journal of neuroendocrinology    June 6, 2008   Volume 20, Issue 8 988-996 doi: 10.1111/j.1365-2826.2008.01751.x
Haritou SJ, Zylstra R, Ralli C, Turner S, Tortonese DJ.Equine pituitary pars intermedia dysfunction (PPID) is a common and serious condition that gives rise to Cushing's disease. In the older horse, it results in hyperadrenocorticism and disrupted energy metabolism, the severity of which varies with the time of year. To gain insight into the mechanism of its pathogenesis, 24-h profiles for peripheral plasma melatonin, serotonin, dopamine and cortisol concentrations were determined at the winter and summer solstices, and the autumn and spring equinoxes in six horses diagnosed with Cushing's disease and six matched controls. The nocturnal rises in p...
Rapid phase adjustment of melatonin and core body temperature rhythms following a 6-h advance of the light/dark cycle in the horse.
Journal of circadian rhythms    August 24, 2007   Volume 5 5 doi: 10.1186/1740-3391-5-5
Murphy BA, Elliott JA, Sessions DR, Vick MM, Kennedy EL, Fitzgerald BP.Rapid displacement across multiple time zones results in a conflict between the new cycle of light and dark and the previously entrained program of the internal circadian clock, a phenomenon known as jet lag. In humans, jet lag is often characterized by malaise, appetite loss, fatigue, disturbed sleep and performance deficit, the consequences of which are of particular concern to athletes hoping to perform optimally at an international destination. As a species renowned for its capacity for athletic performance, the consequences of jet lag are also relevant for the horse. However, the duration...
Variability of plasma melatonin level in pony mares (Equus caballus), comparison with the hybrid: mules and with jennies (Equus asinus).
Reproduction, nutrition, development    December 15, 2006   Volume 46, Issue 6 633-639 doi: 10.1051/rnd:2006043
Guillaume D, Zarazaga LA, Malpaux B, Chemineau P.In long-day breeders like horses, the length of nocturnal melatonin secretion is the main messenger of photoperiod. Previous studies have shown that the nocturnal jugular melatonin concentration is lower in horses, than in mules but is unknown in donkeys. The aim of this study was to estimate the inter-animal variability of plasma melatonin concentration in domestic mares and to compare this concentration with those observed in domestic jennies and in their hybrid mules. In the autumn, blood samples were collected at 22 h, 23 h, 0 h and 1 h during 2 nights at 3 weeks intervals, in 110 pony mar...
Seasonal changes in ovarian activity: lessons learnt from the horse.
Animal reproduction science    December 15, 2006   Volume 100, Issue 3-4 225-242 doi: 10.1016/j.anireprosci.2006.12.001
Donadeu FX, Watson ED.The annual reproductive cycle in the horse involves a reduction in ovarian activity during short days. The absence of ovulatory activity during winter has important consequences for an equine industry eager to breed mares early during the year. The anovulatory season results from a reduction in the secretion of pituitary gonadotropin that is in turn triggered by the inhibitory effects of short photoperiod on the hypothalamus-pituitary axis. Recent studies have provided evidence that the response of the ovaries to endocrine stimuli during the anovulatory season is affected not only by circulati...
Onset of puberty and duration of seasonal cyclicity in Lipizzan fillies.
Equine veterinary journal    July 27, 2006   Volume 38, Issue 4 350-353 doi: 10.2746/042516406777749137
Cebulj-Kadunc N, Cestnik V, Kosec M.The onset of puberty and duration of seasonal cyclicity in Lipizzan fillies and young mares have been determined mostly by empirical data, but very little is known about the hormonal changes at puberty and during the seasonal cyclicity in this breed. Objective: To determine the onset of puberty and the course of the cyclicity in young Lipizzan mares from age 1-4 years. Methods: Blood samples were collected twice a month for 2 years from 46 Lipizzan fillies age 1-4 years. Progesterone concentrations, indicating ovarian activity, were measured using an EIA kit. Values above 6.4 nmol/l were taken...
The pituitary and testicular responses to GnRH challenge between 4 and 14 months of age in thoroughbred colts born in spring and autumn.
Animal reproduction science    September 7, 2005   Volume 88, Issue 3-4 287-298 doi: 10.1016/j.anireprosci.2004.12.008
Brown-Douglas CG, Firth EC, Parkinson TJ, Fennessy PF.Gonadotropin releasing-hormone analogue (buserelin) challenges were carried out every 8 weeks from 4 to 14 months of age on thoroughbred colts born in the spring (n = 6) or autumn (n = 5) to define the onset of puberty. In all colts, luteinizing hormone (LH) secretion followed a seasonal pattern, with high baseline and maximal concentrations in the spring and summer and low concentrations in the winter. Testosterone concentrations were undetectable before spring and, thus, autumn-born colts were younger than spring-born colts when a testosterone response to buserelin was first observed. Mean w...
Effects of short-term feed deprivation and melatonin implants on circadian patterns of leptin in the horse.
Journal of animal science    April 14, 2005   Volume 83, Issue 5 1023-1032 doi: 10.2527/2005.8351023x
Buff PR, Morrison CD, Ganjam VK, Keisler DH.Leptin is a protein hormone produced by adipose tissue that influences hypothalamic mechanisms regulating appetite and energy balance. In species tested thus far, including horses, concentrations of leptin increase as animal fat mass increases. The variables and mechanisms that influence the secretion of leptin are not well known, nor is it known in equine species how the secretion of leptin is influenced by acute alterations in energy balance, circadian patterns, and/or reproductive competence. Our objectives were to determine in horses: 1) whether plasma concentrations of leptin are secreted...
Onset of puberty in pasture-raised Thoroughbreds born in southern hemisphere spring and autumn.
Equine veterinary journal    October 6, 2004   Volume 36, Issue 6 499-504 doi: 10.2746/0425164044877422
Brown-Douglas CG, Firth EC, Parkinson TJ, Fennessy PF.There is little information on age, weight and time of year of puberty in Thoroughbred horses, and the interpretation of such data is difficult due to the wide variety of descriptions of the onset of puberty. Objective: To examine the age, bodyweight and date of onset of puberty in Thoroughbreds born in spring and autumn. Methods: Bodyweight data and blood samples were collected in 59 pasture-raised Thoroughbred horses. Five autumn-born and 18 spring-born colts and 3 autumn-born and 33 spring-born fillies were examined from birth to age 13 and 17 months. A testosterone concentration >2 s.d. ab...
Induced lactation with a dopamine antagonist in mares: different responses between ovariectomized and intact mares.
Reproduction in domestic animals = Zuchthygiene    September 3, 2003   Volume 38, Issue 5 394-400 doi: 10.1046/j.1439-0531.2003.00454.x
Guillaume D, Chavatte-Palmer P, Combarnous Y, Duchamp G, Martinat N, Nagy P, Daels PF.The aim of this study was to compare the effects of treatment with repeated injections of sulpiride (a dopamine D2 antagonist) on prolactin secretion and induced lactation in ovariectomized and intact adult mares and to verify if this induction was possible at the beginning and at the end of the birth season. Two experiments were carried out in September [experiment (expt) 1], and in March (expt 2), in France (48 degrees N). In expt 1, three groups of five mares were tested: intact-control, intact-treated and ovariectomized-treated mares. In expt 2, mares previously subjected to artificial pho...
Potential signals mediating the maintenance of reproductive activity during the non-breeding season of the mare.
Reproduction (Cambridge, England). Supplement    April 18, 2003   Volume 59 115-129 
Fitzgerald BP, Reedy SE, Sessions DR, Powell DM, McManus CJ.The seasonal nature of reproductive activity in mares is widely accepted and considerable attention has focused on the mechanisms that lead to the initiation of the breeding season. In contrast, considerably less information is available about the termination of the breeding season. It is interesting to note that each winter a sub-population of mares continues to undergo oestrous cyclicity during the non-breeding season. Continuation of reproductive activity during the winter occurs most frequently in mares that maintain a non-pregnant condition in successive years. The maintenance of a non-pr...
Photopigment basis for dichromatic color vision in the horse.
Journal of vision    April 8, 2003   Volume 1, Issue 2 80-87 doi: 10.1167/1.2.2
Carroll J, Murphy CJ, Neitz M, Hoeve JN, Neitz J.Horses, like other ungulates, are active in the day, at dusk, dawn, and night; and, they have eyes designed to have both high sensitivity for vision in dim light and good visual acuity under higher light levels (Walls, 1942). Typically, daytime activity is associated with the presence of multiple cone classes and color-vision capacity (Jacobs, 1993). Previous studies in other ungulates, such as pigs, goats, cows, sheep and deer, have shown that they have two spectrally different cone types, and hence, at least the photopigment basis for dichromatic color vision (Neitz & Jacobs, 1989; Jacob...
Behavior of various mammalian albumins towards bilirubin binding and photochemical properties of different bilirubin-albumin complexes.
International journal of biological macromolecules    February 6, 2003   Volume 31, Issue 4-5 187-193 doi: 10.1016/s0141-8130(02)00081-8
Tayyab S, Khan NJ, Khan MA, Kumar Y.Bilirubin (BR) binding properties of serum albumins from different mammalian species viz. human (HSA), equine (ESA), dog (DSA) and guinea pig (GPSA) were studied by absorption, fluorescence and CD spectroscopy. Whereas, a complex of BR with ESA produced maximum change, GPSA-BR complex showed weaker interaction as reflected from absorption and fluorescence spectroscopic data. Conformational analysis of these albumins by near- and far-UV CD spectra suggested similar structural characteristics (both secondary and tertiary structures) for ESA and HSA, whereas, DSA and GPSA had lower amounts of sec...
Unusual ovarian activity in a mare preceding the development of an ovarian granulosa cell tumour.
Australian veterinary journal    August 16, 2002   Volume 80, Issue 1-2 32-36 
Chopin JB, Chopin LK, Knott LM, de Kretser DM, Dowsett KF.An 8-year-old mare, with a foal at foot, was inseminated on foal heat with frozen semen, with the resultant pregnancy lost between days 34 and 41. The right ovary developed a large anovulatory follicle that was non-responsive to multiple doses of ovulating agents. The follicle eventually appeared to luteinise, although plasma progesterone concentrations did not reflect this. Another follicle developed, responded to GnRH and resulted in a pregnancy from frozen semen that went to term with a healthy foal. When the mare was examined after foaling, the structure on the right ovary appeared to be a...