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Reproduction The Mammalian Strategy: • • • Relatively few intrauterine young (higher survival rate) Nourish neonates with milk (high survival early; bonding) Young remains with mother (or parents) at minimum until weaned (parental protection; learned behaviors) Reproduction The Mammalian Strategy: • • Amount of energy invested per young is lower than nonmammals; Relatively few young produced but most survive to potentially reproduce Costs of Lactation K c a l/d a y /in d iv id u a l C a lo ric In ta k e o f B a n k V o le s 50 40 B re e d in g fe m a le 30 20 N o n -b re e d in g fe m a le 10 S o u rc e : F lo w e rd e w (1 9 8 7 , M a m m a ls : th e ir re p ro d u c tiv e b io lo g y a n d p o p u la tio n e c o lo g y ) 0 0 18 P re g n a n c y 36 L a c ta tio n M u ltip le s o f M a in te n a n c e Tradeoffs in Litter Size L a c t a t io n C o s t s f o r C a t s 4 4 5 K it t e n s 3 3 3 K it t e n s 2 2 2 K it t e n s 1 1 M a in t e n a n c e n e e d s 0 0 0 1 2 3 W e e k o f L a c t a t io n 4 5 Reproductive Endocrinology “Crash Course” * Feedback mechanisms (environmental stimuli; hormone secretions) Reproductive Endocrinology “Crash Course” Ovarian Cycle Influenced by: 1) Follicle stimulating hormone (FSH) and luteinizing hormone (LH) secreted by pituitary • follicle growth which triggers ovary to secrete estrogen Reproductive Endocrinology “Crash Course” Ovarian Cycle Influenced by: 2) Estrogen secretion feeds-back to hypothalamus-pituitary; more LH secreted & less FSH • Ovulation & corpus luteum formation (spongy body which forms in place of ruptured follicle) • Corpus luteum secretes progesterone for uterine wall preparation Reproductive Endocrinology “Crash Course” Ovarian Cycle Influenced by: 3) No fertilization • Corpus luteum recedes to Corpus albicans • Progesterone & estrogen level drop • Begin again in cycle Reproductive Endocrinology “Crash Course” Ovarian Cycle Influenced by: 3) If fertilization occurs… • Corpus luteum continues to produce progesterone for maintaining pregnancy • Placenta soon assumes estrogen & progesterone secretion amnion chorion embryo allantois Four Major Parts of Embryonic Membranes 1) yolk sac: part of primitive intestine lying external to embryo; forms from endoderm • No nutritional value • Portion of placenta in some cases (e.g., marsupials) Four Major Parts of Embryonic Membranes 2) amnion: forms from ectoderm & mesoderm around the embryo • Filled with serous fluid = prevent dessication/shock 3) allantois: out-pocket from hindgut of embryo • Movement of nutrients & O2 • Forms blood vessels for placenta Four Major Parts of Embryonic Membranes 4) chorion: outer embryonic layer (ectoderm); envelopes entire assemblage • villi • contact with uterine wall placenta: includes embryonic membranes & lining of uterine wall (endometerium) Types of Placenta A) Placenta types based on villi distribution on chorion: 1) diffuse: villi scattered over entire surface of chorion = increased SA for absorption e.g., lemurs, perissodactyls, some artiodactyls 2) polycotyledonary: islands of villi scattered over chorion e.g., other artiodactyls such as bovids Types of Placenta A) Placenta types based on villi distribution on chorion: 3) zonary: band of villi encircle center of blastocyst; lacking villi elsewhere e.g., carnivores 4) discoidal: regional restriction of villi e.g., most mammals discoidal zonary diffuse Types of Placenta B) Placenta type based on connection between villi & endometrium: 1) nondeciduate: loose fitting of villi with endometrium; villi pull free without disrupting endometrium during parturition (whales, ungulates) 2) deciduate: close fitting of villi-endometrium; villi pull free & cause erosion of endometrium during parturition (rodents, carnivores) Types of Placenta C) Placenta type based on degree of intimacy between embryonic & maternal parts: 1) choriovitelline: blastocyst lies in endometrium depression; does not embed 2) chorioallantoic: villi; blastocyst rests against endometrium at allantoischorion contact point Types of Placenta C) Chorioallantoic Placenta Types: 1) epitheliochorial – lemurs, cetaceans, equids, suids - epithelial cells of chorion in contact with epithelial cells of uterus; villi in pockets in endometrium 2) syndesmochorial – artiodactyls - lacking uterine epithelial barrier; contact uterine tissue Types of Placenta C) Chorioallantoic Placenta Types: 3) endotheliochorial – carnivores - epithelial cells of chorion in contact lining of uterine capillaries 4) hemochorial – insectivores, bats, higher primates - villi in direct contact with maternal blood Types of Placenta C) Chorioallantoic Placenta Types: 5) hemoendothelial – lagomorphs, some rodents - lining of villi blood vessels only barrier to maternal blood Reproductive Patterns 1) Continuous embryonic development (“typical”) a) ova fertilized in oviduct b) zygote begins mitosis - descends towards uterus c) zygote reaches uterus – mitosis ongoing – reaches blastocyst stage as implanting into endometrium d) placental connection: uterus to embryo e) continual development until parturition Reproductive Physiology - Implantation of embryo in uterine wall for varying lengths of time - Embryo supplied with nutrients via the placenta Reproductive Patterns 2) Deviations from contiuous development strategy: a) Delayed Fertilization: ovulation & fertilization delayed until an extended time after copulation • • • Viable sperm retained in female Ovulation occurs ~months after copulation Common to many temperate bats (vespertilionids) Reproductive Patterns Example Fall copulation Winter Sperm storage 2) Deviations from contiuous development strategy: a) Delayed Fertilization: Early spring ovulation Spring-summer Embryo develops after fertilization Reproductive Patterns 2) Deviations from contiuous development strategy: b) Delayed Development: blastocyst embeds into endometrium & then becomes dormant; development delayed (e.g., bats) Reproductive Patterns 2) Deviations from contiuous development strategy: Example Late summer Summer-Fall Blastocyst forms Blastocyst dormant Late fallearly winter Development begins Early spring parturition b) Delayed Development: Reproductive Patterns 2) Deviations from contiuous development strategy: c) Delayed Implantation: obligate & facultative examples e.g., weasels, seals, bears • Blastocyst forms but does not embed & ceases to develop • Floating blastocyst remains dormant 2 weeks to 1 year Reproductive Patterns Summer Mating (Jun-Jul) 2004 March 2005 Implantation (8-9 mo delay) Spring Parturition (Apr-May) 2005 Summer Mating (Jun-Jul) (including 2005 2005 females 2) Deviations from contiuous development strategy: c) Delayed Implantation: e.g., Mustela erminea (avg age at death = 1.5 to 2 yrs) *gestation period = 9-10 months Reproductive Patterns Spring-Summer (Apr-May) 2004 Spring-Summer (May-Jun) 2004 Summer (Jul-Aug) 2004 Summer-Fall Aug-Sep 2004 Mating Parturition Mating? Sexually Mature 2004 Females Parturition (2nd litter) Mustela nivalis Delayed Implantation???? * NO (avg age at death = <1 yrs) * gestation period = 3537 days • 2 litter per year possible • Relation to vole cycles Types of Breeding Seasons 1) Continuous – year round breeding; no seasonality; common to tropics 2) Restricted Optimal timing for: a) Regular – seasonal breeding; temperate regions b) Irregular – discontiuous breeding during rainfall, etc… desert/arid regions * mating (time with best availability of mates) * birth (time with abundant resources Seasonality to Mating & Parturition based on resource availability (i.e, mates or food) Resources Gestation Period Mating Fall Birthing Winter Spring Summer Body size relation to length of gestation period….What if mammal could “extend” the gestation period to birth in a more favorable time and/or insure mating opportunities? (e.g., weasels) Gestation Period Resources Delay Mating Fall Major Development Birthing Winter Spring Summer Reproduction Sexual Maturity (puberty) – age when capable of producing gametes influence onset/cessation (restricted) *environmental factors efficiency of reproduction (continuous) Influences on Puberty & Reproduction 1) Light (photoperiod) Rattus norvegicus (continuous breeder) • • • normal light continuous light = 6 days earlier than normal (FSH) Constant dark = 16 days later than cont. light Influences on Puberty & Reproduction 1) Light (photoperiod) Microtus arvalis (seasonal breeder) • • 1) 2) 3) 4) 5) breeds 21 Mar – 24 Jun simulate photoperiod during (22 Sep – Dec) Natural light Artificial light Uniform 16-h daylength Uniform 8-h daylength until Nov, then 13-h day Control (“out of season”) Results…. • #1-4 = reached puberty • >60% females = pregnant • Control = no reproduction/puberty *Light (photoperiod) linked to reproductive development Influences on Puberty & Reproduction 2) Temperature rodents Temp Puberty 1st Estrus Experimental Animals -3oC 33 days 61 days Control 21oC 26 days 38 days **Growth rates lowered due indirectly to low temps. Thus, results directly in delayed puberty Influences on Puberty & Reproduction 3) Nutrition – under-nutrition delays puberty in both females and males 4) Precipitation – deer in Texas (Knowlton) - “high” rainfall lead to shorter breeding season, more synchronous breeding & fawning - lower predation rates (functional response of coyotes) # prey consumed Prey density Influences on Puberty & Reproduction 5) Social Effects/Density (examples from captive mice) Lee-Boot Effect: pseudopregnancy induced among crowded females; may go anestrus Whitten Effect: synchronized estrus cycles when male introduced into population of females Bruce Effect: implantation blocked, pregnancy aborted if females exposed to strange, new male * Male urine stimulates FSH & LH secretion (pheromones) Readings • Reproductive Cycles & Life-History Strategies, pp. 354-356 • Litter Size & Reproductive “Seasons”, pp. 356-357 • Lactation and Postnatal Growth, pp. 359-363