Characterization of the estrous cycle in female beluga ( Delphinapterus leucas ) using urinary endocrine monitoring and transabdominal ultrasound: Evidence of facultative induced ovulation

Highlights ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are facultative induced ovulators with evidence of spontaneous ovulation.

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Published in:	General and comparative endocrinology Vol. 175; no. 3; pp. 389 - 397
Main Authors:	Steinman, K.J, O’Brien, J.K, Monfort, S.L, Robeck, T.R
Format:	Journal Article
Language:	English
Published:	United States Elsevier Inc 01-02-2012
Subjects:	adults animal ovaries Animals assisted reproductive technologies Beluga Whale - physiology conjugated estrogens Delphinapterus leucas Endocrinology & Metabolism Estrogens - urine Estrous cycle Estrous Cycle - physiology excretion Female Follicular ultrasonography Gonadal Steroid Hormones - urine Immunoassay immunoassays Induced ovulation luteinizing hormone Luteinizing Hormone - urine Male monitoring Ovary - diagnostic imaging ovulation Ovulation - physiology Progestins - urine Prolactin Prolactin - urine Reproducibility of Results Sexual Behavior, Animal - physiology ultrasonics Ultrasonography White whale Estrous cycle Follicular ultrasonography White whale Prolactin Induced ovulation
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Abstract	Highlights ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are facultative induced ovulators with evidence of spontaneous ovulation.
AbstractList	Recent, successful application of assisted reproductive technologies in captive beluga has resulted from the extensive study of male beluga reproductive biology. Optimization of assisted reproduction requires additional detailed knowledge of the female estrous cycle. Our specific objectives were to: (1) validate urinary immunoassays for use in this species; (2) elucidate annual ovarian cycle dynamics through the combined use of hormone excretion patterns and transabdominal ultrasound; and (3) establish whether ovulation in this species is spontaneous or induced by male factors. Ovulation was observed in four of 15 estrous cycles monitored in four adult female beluga maintained in a single-sex group. After introduction of a breeding male, ovulation was observed in six of seven estrous cycles. All estrous cycles occurred from March through June. For spontaneous ovulations (n=4), the inter-estrous interval was 34d (range 33-35d), with a follicular phase length (FPL) of 25±8d (mean±SD). For all ovulatory estrous cycles (with and without a breeding male), urinary estrogen conjugates (EC, 15.3±7.9ng/mg Cr) and ovulatory luteinizing hormone (ovLH, 17.1±6.6ng/mg Cr) concentrations both peaked on Day 0, and EC concentrations returned to baseline 8±7d later. For non-conceptive cycles, urinary progestagen (Pg) concentrations increased on Day 0 (3.5±1.7ng/mg Cr), peaked on Day+19 (19.7±17.1ng/mg Cr), and were elevated above baseline for 27±4d. Preovulatory follicular diameter and circumference on Day -2±2 (range: Day -4 to -1) from peak EC were 2.5±0.7 and 7.8±1.3cm, respectively. The FPL in non-ovulatory estrous cycles (n=11) lasted 24±10d and EC concentrations gradually declined to baseline over a 21±10d interval following the EC peak (27.8±28.8ng/mg Cr). Non-ovulatory estrous cycles were characterized by the absence of an ovLH surge and no concomitant increase in Pg concentrations above baseline excretion; the mean follicular diameter at or near peak EC was 3.1±0.8cm on Day 2 ±3d from peak EC (range: -1 to +5days from peak EC). Overall, these data confirm that captive beluga exhibit reproductive seasonality and demonstrate that the species is a facultative-induced ovulator. Recent, successful application of assisted reproductive technologies in captive beluga has resulted from the extensive study of male beluga reproductive biology. Optimization of assisted reproduction requires additional detailed knowledge of the female estrous cycle. Our specific objectives were to: (1) validate urinary immunoassays for use in this species; (2) elucidate annual ovarian cycle dynamics through the combined use of hormone excretion patterns and transabdominal ultrasound; and (3) establish whether ovulation in this species is spontaneous or induced by male factors. Ovulation was observed in four of 15 estrous cycles monitored in four adult female beluga maintained in a single-sex group. After introduction of a breeding male, ovulation was observed in six of seven estrous cycles. All estrous cycles occurred from March through June. For spontaneous ovulations (n=4), the inter-estrous interval was 34d (range 33–35d), with a follicular phase length (FPL) of 25±8d (mean±SD). For all ovulatory estrous cycles (with and without a breeding male), urinary estrogen conjugates (EC, 15.3±7.9ng/mg Cr) and ovulatory luteinizing hormone (ovLH, 17.1±6.6ng/mg Cr) concentrations both peaked on Day 0, and EC concentrations returned to baseline 8±7d later. For non-conceptive cycles, urinary progestagen (Pg) concentrations increased on Day 0 (3.5±1.7ng/mg Cr), peaked on Day+19 (19.7±17.1ng/mg Cr), and were elevated above baseline for 27±4d. Preovulatory follicular diameter and circumference on Day −2±2 (range: Day −4 to −1) from peak EC were 2.5±0.7 and 7.8±1.3cm, respectively. The FPL in non-ovulatory estrous cycles (n=11) lasted 24±10d and EC concentrations gradually declined to baseline over a 21±10d interval following the EC peak (27.8±28.8ng/mg Cr). Non-ovulatory estrous cycles were characterized by the absence of an ovLH surge and no concomitant increase in Pg concentrations above baseline excretion; the mean follicular diameter at or near peak EC was 3.1±0.8cm on Day 2 ±3d from peak EC (range: −1 to +5days from peak EC). Overall, these data confirm that captive beluga exhibit reproductive seasonality and demonstrate that the species is a facultative-induced ovulator. ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are facultative induced ovulators with evidence of spontaneous ovulation. Recent, successful application of assisted reproductive technologies in captive beluga has resulted from the extensive study of male beluga reproductive biology. Optimization of assisted reproduction requires additional detailed knowledge of the female estrous cycle. Our specific objectives were to: (1) validate urinary immunoassays for use in this species; (2) elucidate annual ovarian cycle dynamics through the combined use of hormone excretion patterns and transabdominal ultrasound; and (3) establish whether ovulation in this species is spontaneous or induced by male factors. Ovulation was observed in four of 15 estrous cycles monitored in four adult female beluga maintained in a single-sex group. After introduction of a breeding male, ovulation was observed in six of seven estrous cycles. All estrous cycles occurred from March through June. For spontaneous ovulations ( n = 4), the inter-estrous interval was 34 d (range 33–35 d), with a follicular phase length (FPL) of 25 ± 8 d (mean ± SD). For all ovulatory estrous cycles (with and without a breeding male), urinary estrogen conjugates (EC, 15.3 ± 7.9 ng/mg Cr) and ovulatory luteinizing hormone (ovLH, 17.1 ± 6.6 ng/mg Cr) concentrations both peaked on Day 0, and EC concentrations returned to baseline 8 ± 7 d later. For non-conceptive cycles, urinary progestagen (Pg) concentrations increased on Day 0 (3.5 ± 1.7 ng/mg Cr), peaked on Day + 19 (19.7 ± 17.1 ng/mg Cr), and were elevated above baseline for 27 ± 4 d. Preovulatory follicular diameter and circumference on Day −2 ± 2 (range: Day −4 to −1) from peak EC were 2.5 ± 0.7 and 7.8 ± 1.3 cm, respectively. The FPL in non-ovulatory estrous cycles ( n = 11) lasted 24 ± 10 d and EC concentrations gradually declined to baseline over a 21 ± 10 d interval following the EC peak (27.8 ± 28.8 ng/mg Cr). Non-ovulatory estrous cycles were characterized by the absence of an ovLH surge and no concomitant increase in Pg concentrations above baseline excretion; the mean follicular diameter at or near peak EC was 3.1 ± 0.8 cm on Day 2 ±3 d from peak EC (range: −1 to +5 days from peak EC). Overall, these data confirm that captive beluga exhibit reproductive seasonality and demonstrate that the species is a facultative-induced ovulator. Highlights ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are facultative induced ovulators with evidence of spontaneous ovulation.
Author	Robeck, T.R O’Brien, J.K Steinman, K.J Monfort, S.L
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Keywords	Estrous cycle Follicular ultrasonography White whale Prolactin Induced ovulation
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Snippet	Highlights ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are... ► Urinary hormone enzyme immunoassays and ovarian ultrasound validated in the beluga. ► Beluga demonstrate reproductive seasonality. ► Belugas are facultative... Recent, successful application of assisted reproductive technologies in captive beluga has resulted from the extensive study of male beluga reproductive...
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SubjectTerms	adults animal ovaries Animals assisted reproductive technologies Beluga Whale - physiology conjugated estrogens Delphinapterus leucas Endocrinology & Metabolism Estrogens - urine Estrous cycle Estrous Cycle - physiology excretion Female Follicular ultrasonography Gonadal Steroid Hormones - urine Immunoassay immunoassays Induced ovulation luteinizing hormone Luteinizing Hormone - urine Male monitoring Ovary - diagnostic imaging ovulation Ovulation - physiology Progestins - urine Prolactin Prolactin - urine Reproducibility of Results Sexual Behavior, Animal - physiology ultrasonics Ultrasonography White whale
Title	Characterization of the estrous cycle in female beluga ( Delphinapterus leucas ) using urinary endocrine monitoring and transabdominal ultrasound: Evidence of facultative induced ovulation
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