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The Effect of Photoperiod on the Gonads and Basophilic Adenohypophyseal Cells of the Rough-Skinned Newt, Taricha Granulosa Granulosa

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The Effect of Photoperiod on the Gonads and Basophilic Adenohypophyseal Cells of the Rough-Skinned Newt, Taricha Granulosa Granulosa University of the Pacific Scholarly Commons University of the Pacific Theses and Dissertations Graduate School 1970 The effect of photoperiod on the gonads and basophilic adenohypophyseal cells of the rough-skinned newt, Taricha granulosa granulosa Paul Patton Coambs University of the Pacific Follow this and additional works at: https://scholarlycommons.pacific.edu/uop_etds Part of the Biology Commons Recommended Citation Coambs, Paul Patton. (1970). The effect of photoperiod on the gonads and basophilic adenohypophyseal cells of the rough-skinned newt, Taricha granulosa granulosa. University of the Pacific, Thesis. https://scholarlycommons.pacific.edu/uop_etds/1714 This Thesis is brought to you for free and open access by the Graduate School at Scholarly Commons. It has been accepted for inclusion in University of the Pacific Theses and Dissertations by an authorized administrator of Scholarly Commons. For more information, please contact [email protected]. ~HE EFFECT OF PHOTOPERIOD ON THE GONADS ~----- AND BASOPHILIC ADENORYPOPHYSEAL CELLS OF THE ROUGH-SKINNED NEWT, TARICH.A GRANULOSA GR.tU'fULOS.A ---·--- A Thesis Presented to the Faculty of the Department of Biological Sciences University of the Pacific In Partial Fulfillment, of the Requirements for the Degree Mester of Scie:nc.e in Biology by Paul Patton Coambs August 1970 This thesis, written and submitted by Paul Patton Coambs is approved for recommendation to the Graduate Council, University of the Pacific. 1besis Committee: Acknowledgements 'The author wishes to express his app1•ecia·t1on. to Drs. Lee Christianson, Anne Mathias and M. Dale Arvey, all of the University of the Pacific, for help and direction of this study, Grateful acknowledgement is made to !4r. Arnold Coa.mbs for his technical assistantship in building the st1.tdy area. The author is especially indebted to his wife, Dianne Coambs, for enoourgement and invaluable assistantship during the entire study. 111 Table of Contents Page c List of Tables • • • • • • • • • • • • • • • • • • • • • v List of Fig-..1res • • • • • • • • • • • • • • • • • • • • v1 Chapter 1. Introduction • • • • • • • • • • • • • • • .. • • • 1 2. lJatural History • • • • • • • • • • • • • • • • • 4 ). Materials and Methods • • • • • • • • • • • • • • 6 4. Results • • • • • • • • • • • • • • • • • • • • • 9 s. Discussion • • • • • • • • • • • • • • • • • • • • 28 6. Summary • • • • • • • • • • • • • • • • • • • • • :n Literature Cited • • • • • • • • • • • • • • • • • • • • 34 iv .. ----- - ~ List of Tables _ l~ Table Page 1. Data for Taricha g_. _granulosa Subjected to 24. - - ------ Hour Light Photoperiod • • • • , • • • • • • • • 11 2. Data for Iaricha £• granule~ Subjected to 12 Hour Light-12 Hour Dark Photoperiod. , • • • • • 1:3 J, Data for Taricha g. g_t.._anulosa Subjecte.d to 214- Hour Dark Photoperiod • • • • • • • • • • • • • 15 = E~-------- v List of Fi.gures Figure Page 1. The Range and Median Percent of Water in the Gonads in Both Mlae and Female Taricha IS.• aramllosa Subjected to 24 HourLight. Photoperiod • . • • • • • • • • • • • • • o • • • 17 2. The Range an.d I1edian Percent of Water. in the Gonads in Both Male and Female 'I'ar!.cha S.• e;~~ulosa Subjected to 12 Hour Light-12 Hour Dark.Photoperlod •••••• , ••• • • • 18 ). The Range and 11ed1an Percent of Wate.r in the Gonads in Both Male and Female Taricha IS.• B!anulosa Subjected to.24 Hour Light Photoperiod • ·,., _. , ,"' . • • • • • • ., • • • • • 19 4. The J.I<Iedlan Percentage of Water in the Testes of ·'l'a_!icha g. £l.!:&nulosa under the .E:nvj.ron­ mental Ccndi ti(>nS of '2.4 Hour Light, 12 Hour Ligh.t-12 Hour Dark or 24 Hour· Dark Photo·· period • • • ~ • • • • • •. • _, • • . • • • • • • 20 .s. The llfed:!.an and Range of W0t Gonad Weights of Both Hale and Female Tar1cha g. granulosa Subjected to 24 Hour Light Photoperiod • • • • • 21 6. The Median and Range of Wet Gonad Weights of Both Male and Female Tarich~~· JEanulosa Subjected to 12 Hour Light-12 Eour Dark e.-- PhotoperiOd • • e • • • • • • • • • • • ~ • • • 22 ?. The Median and Range of Wet Gonad Weights o.f Both Male and Female Taricha e;. granulosa Subjected to 24 Hour Dark Photoperiod • • • • • 2) a. The ~'iedian Wet Testes \~eight for Tarioha s_. granulosa Subjected to each Environmental Condition • • . a • • • • • • • • • • • • • • • • 24 The Median Wet Ovary Weight for Taricha g, gr~plos~ Subjected to.eaoh EnVironmental Condition ................. • • • 25 10. The Range and Medirul Basophilic Adenohypophyseal Cell Number Subject to each Environmental Cond1 t1 on • .. .. • • ., • • • • • • • • • • • • • 26 Vi v11 F1gure 11. The .l~edian Basophilic Adenohypo;physeal Cell Number Subject to each Environmental Cond1 tion • • • • • • • • • • ~ • • • • • a • • 27 Chapter. 1. -=- -------- ---- ~---- Introduction A variety of factors, temperature, photoperiod, rainfall, food availability and others, have been found to stinr.1late reproductive and migrational activity in various vertebrates. Seasonal variation in light has been well studied as the causative factor in the stimulation of reproductive and migrational activity in fish. birds, and mammals. Light is generalYy believed to be the most critical 'single environmental factor in regulating the development and periodicity of reproductive activity (Gorbm.an and Bern, 1962). · Sexual periodicity is not a direct function of the external environment but is governed by two main factors, internal gonadal rhythm and an external environmental stimuli. The former is characteristic of the species while the latter -- - ----- determines the precise time of breeding (Bullough, 1951). --- Both Internal and external factors have been shown to play a part in the annual testicular cycle in the frog ~ temporaria. However, in this situation, it is temperature and not photoperiod which is the important external factor (van Oordt, et al., 1959). In the frogs Rana esculenta, R. sraeca and Leptodactylus ocellatus, in the newts Tritu.rus cristatus carnifex and T. viri.descens and in the 1 2 •salamander Geotriton fuscus, tempe:t·atures of approximately . '4' 20 degrees centigrade induced spermatogenesis while lower temperatures inhibited it. Telmatobius schreiteri and Erla raddiana andina which live in a:r.:eas of low winter temperature and the tropical and subtropice.l species, ~ erytrea, E,. gr~, R. tigrina, ].!:!!2, melan.ostictus, 1!. arenarum, ,£. paracnemis, ;g. granulos1lll. d'orbi.r,nY.b Leptodactylus ocellatus reticulata, 1· 2.• bonaerensis, have continuous spermatogenesis {van Tienhoven, 1968). ~erefore, spermatogenesis may or may not be independent of temperature. Bainf'all appears to be.another stimulator of reproductive activity, especially the reproductive :nigration prior to breeding {Gorbman and Bern, 1962) •. Packer {1960) has shown that rainfall is the important factor in ini tiati:rig' the reproductive·migration of Taricha rivularis. It has been sho~~ that a wide variety of vertebrates have become adapted such that increasing photoperiod acts as --- an 1n1 tiator for the reproductive activity.. It is . equally The 'function of light in the reproductive cycle in amph1 bians has not been extensively studied. However·, Farner ( 1965) feels that the small amount of experimental evidence suggests that photoperiod may play a role in the testicular cycle of ~ esculenta. The purpose of this paper is to study the effect of light on the gonads and the gonadotropic hormone forming cells of the adenohypophysis in the newt "-'----=---=-----·-· Taricha granulosa granulosa. ---- ! ' I ...; _______ -- -- --- - '-=-------- Chapter 2 :.0:---~-~ Natural History Recent taxonomic treatments on the genus Taricha, family Salamandridae, have shown a difference of opinion on the relationships of the group. Riemer (1958) recognizes three species, I· rin1laris, I· torosa and !• granulosa, and upholds the subspecies of I· !· slerrae and I· ~· mazama~. Pimentel {1958, 1959), however, recognizes two species, I• r1vular1s and I· torosa, and proposes that the six subspecies I• !· torosa, I· !· sierra~, I· S• granulosa, I· ~· ~azamae, I• li• tw1tty1 and I· g. !timulans be assigned to the species ~. torosa. Stebbins (1966) lists three species of Tar~ ' and has therefore apparently agreed with Riemer• s classlf1cation. While the author must admit that he has observed animals which would support combining T. granulosa and I· torosa in the same species, he will submit to the authority of Stebbins. The natural history of members of the. genus Taricha is well documented (Ritter, 1897; Chandler, 1918; Storer, 1925; Smith, 1941; Twitty, 1942; Stebbins, 1951, 1951l-, 1966; M1.ller and Robbins, 1954; Riemer, 1958; Packer, 1960; Pimentel, 1960; and Storer and Usinger, 1966), T. granulosa, the rough-skinned newt, has the most extensive range and 1s the most aquatic of the three species 4 of Tarich~. Stebbins .(1954. 1966) gives the range of !• £~Ul~ as extending along.the coast from southeast .Alaska to south Santa Cruz County, California, . west of. the o------~·' Cascade Mountains, and south in the foothills of the Sierra ~ : Nevada to Magalia., Butte County, California. There is also an isolated segment of the species which extends from the Moscow Mountains. I,ata.h County, Idaho, to Thompson Falls, Sanders County • Monta:'la, :r_. li• mazamae is found around Crater Lake, Crater Lake National Park, Oregon, and T. a. gran~losa inhabits the rest of the range of the species. The newt can ·be found in humid coastal i'orests, grasslands, ponds, reservoirs and slo.w moving streams. 1:_. gran'::!Josa,
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