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EinleitungMit Beginn seiner Domestikation werden die Lebensbedingungen desRindes nicht mehr in erster Linie durch natnliche Umwelteinflssebestimmt, sondern in zunehmendem Mae von den besonderenUmweltbedingungen, denen der Mensch unterliegt, die er sich und denTieren schafft. Das Rind wird seitdem durch die sich entwickelndenMglichkeiten und die sich verndernden Ansprche der Mensehengeprgt. So war der Beginn des Ackerbaus Voraussetztung fr dieDomestika[ion des Rindes; gleichzeitig wurde mit ihm dieArbeitskraft der Tiere erschlossen. Im Laufe der Geschichte erlangtdas Rind neben seiner zen[ralen Aufgabe als Arbeitstier je nachRegion und Kultur Bedeutung als Dung- und Rohstofflieferant, alsOpfertier oder Tauschobjekt, und immer strker auch alsNahrungslieferant. In Mitteleuropa war bis Mitte des 15. Jahrhundertsin Gebieten mit gutem natrlichen Futterwuchs die Ochsenmast diewichtigste Einnahmequelle aus der Rinderhaltung, anschlieendentwickelte sich mit dem Anwachsen der Stdte dieMilchviehhaltung. Mit der systematischen Einflunahme desMenschen auf die Rinderzucht begann Mitte des lB. Jahrhunderts diePeriode der Rassenbildung und ein Verdrngungsproze deranspruchslosen Landschlge durch leistungsfhige Rassen aus denfutterwchsigen Grnlandgebieten (IB).Die Anwendung des wissenschaftlichen und technischen Fontschrittskennzeichnet die rasante Entwicklunq der landwirtschaf tlichenProduktion des 20. Jahrhunderts. Arbeitswirtschaf tliche,technologische und konomische Aspekte treiben die Spezialisierungder Produktionsrichtungen und die Intensivierung der Tierproduktionvoran. Mit der organisierten und systematischen Selektion aufbestimmte Leistungen innerhalb von Populationen pa0t sich dieTierzuchl an die s[eigenden Ansprche an. 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Diese als "biotechnisch'lbezeichneten Manahmen finden ihre Anwendung v.a. im Bereich derReproduktion , d.h. sie gneifen in die das Fortpflanzungsgeschehensteuernden Mechanismen ein, um sie in gewnschtem Sinne zeitlichoder quantitativ zu beeinflussen.Eine der bedeutensten biotechnischen Manahmen ist die knstlicheBesamung, die weltweit in der Tierzucht angewandt wird undentscheidend zum Zuchtfortschritt beitrgt. Weitere bioLechnischeEingriffe sind z.B. der Embryotransfer, oder aber der Einsatz vonHormonen zur Steuerung des Geschlechtszyklus weiblicher Haustiere. Eine der ltesten biotechnischen Manahmen ist die Kastration. Beimmnnlichen Mastschwein wird sie routinemig durchgefhrt, um denproduktionsschdigenden Geschlechtsgeruch zu vermeiden. Beim Stierwurde sie in der Vergangenheit angewandtr urrr sein aggressivesVerhalten auszuschalten und die Ochsen so als Arbeitstiereinzusetzten oder um unter extensiven Bedingungen Weidemast zuermglichen.Heute wird die tierische Arbeitskraft durch Maschinen ersetzl. Unterden intensiven Fut.terverhltnissen in Mitteleuropa haL sich dieStallmast von Jungbullen weitestgehend durchgesetzt. Neben dergenerellen Entwicklung der Landwirtschaft hat dazu auch dasvernderte Bedrfnis des Verbrauchers nach magefem Fleischbeigetragen. In Lndern mit weiten extensiven Weideflchen, wie inUSA und Argentinien, wird der Nachteil der um ca. 15 % geringerenZunahmen in der Ochsenmast durch den Einsatz von synthetischenHormonen aufgehoben. 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Bei Anwendung inanabole Wirkungen zu erwarten,auen zugefhrter Hormonebeeintrchtigen (15). der Bullenmast sind deshalb kaumim Gegenteil, eine berdosis vonkann den Mastefekt sogar Mit der Entwicklung von der traditionellen Anbindehaltung hin zurwirtschaf tlicheren Laufstallhaltung bleibt die Suche nachbiotechnischen Manahmen auch in der Bullenmast aktuell: Die inGruppen gehaltenen Tiere knnen ihne soziale Aktivitt slrkerentf alten; die Folge sind zunahmemindernde Unruhe sowie dieVerletzungsrate erhhende Kmpfe und damit wirtschaf tlicheEinbuen. Mit den bisher eingesetzten, meist antiandrogen wirkendenMitteln konnte zwar eine sexuelle Ruhigstellung erreicht werden, siebeeintrchtigten jedoch gleichzeitig die anabo.le Wirkung derkrpereigenen Testosteronproduktion, was teilweise eineVerminderung der tglichen Zunahmen zur Folge hat. Man forschtdeshalb weiterhin nach Mglichkeiten, die Verhaltensaktivitt derStiere zu hemmen, ohne gleichzeitig die anabole Androgenpotenz zumindern, wenn mglich diese sogar zu frdern. Dabei geht man vonder Annahme aus, da das test,osteronabhngige Verhaltenempfindlicher und leichter zu blockieren ist, als dietestosteronabhngigen Stoffwechsel- und Organeffekte (5).Zwei grundstzliche Probleme ergeben sich bei Applikation vonHormonen zu Mastzwecken: Die Tiere reagieren individuelluntersehiedlich auf eine bestimmte Dosis. Auerdem ergibt sich v.a.bei krperfremden Verbindungen die vielschichtige Problematik derRckstandsbildung; sie hngt sowohl von exogenen Faktoren, wie vonder Applikationsart, als auch von endogenen Faktoren, wie derVerstoffwechselungsrate ab. KARG unterscheidet in einem ArtikelrrNutzen und Risiko der Hormonmast" zwischen "hantenrr und t'relativharmlosen" Hormonen. Zu letzt,eren zhlen z.B. die natrlichenSexualhormone wie stradiol, Testosteron und Progesteron. Zu den"hanten'r strogenhaltigen Anabolika zhlen Stilbenderivate wieDithylstilbstrol (DES) (f 6). 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-5 2. ProblemstellungEine biotechnische Manahme, mit deren Auswirkungen sich dieseArbeit beschftigL, ist der knstlich herbeigefhrte Knyptorchismus.Hiermit wird in die endokrinen Regelmechanismen eingegrif f en,jedoch ohne uenen Einsatz von Sexagenverbindungen. Diekrpereigene Hormonproduktion wird durch Eingrif f in dieTestisfunktion beeinflut, und damit auch die sekundrenSexualmerkmale Verhalten und Gewichtsentwicklung.In Australien wurde bereits Ende der 50er Jahre der knstlicheKryptorchismus mit Versuchen in der Lmmermast untersucht:Unkastrierte Schafbcke haben im Vergleich zum Hammel ca. 15 o/ohhere Zunahmen, ihr Fleisch wird jedoch vom Verbraucher wegender geschmacklichen. Unterschiede nicht akzeptiert. Diekryptorchiden Tiere 'erreichten die Zunahmen der unbehandeltenLmmer, bei den sensorischen Eigenschaften waren im Vergleich zuHammelfleisch keine Unterschiede feststellbar (4).Ahnliche Ergebnisse erzielten 1971 GLIMP et al. in Versuehen mitStieren der Rassen Angus und Hereford. Weitere in USAdurchgefhrte Experimente besttigen die Annahme, da die alsI'short scrotum bulls" (ss bulls) bezeichneten Kryptorchiden sowohl inihren Zunahmen, als auch in Schlachtknperzusammensetzung undFleischqualitt mit den LeisIungen der Stiere vergleichbar sind(tau. 1) (l).In einem anderen mit HF-Bullen durchgefhrten Versuch ergibt sicheine Uberlegenheit der Kryptorchiden gegenber den Ochsen; in denZunahmen sind die Stienen den beiden anderen Vergleichsgruppenberlegen (8). Dies tt vermuten, da rassenbedingte Unterschiedein der Reaktion auf den knstlichen Kryptorchismus auftreten.Insgesamt haben jedoch die Ergebnisse der in USA dunchgefhrten endstream endobj 73 0 obj 1699 endobj 74 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 75 0 obj << /Length 45 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj69 Do Q endstream endobj 76 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj69 69 0 R >> /Subtype 72 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 74 0 R 75 0 R 70 0 R ] >> endobj 77 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 37400 >> stream &?*.k>@tp*)5|᥃ԊG_mp8aV(?ieLpTҥ]jZEGW4&?%`u+Ӣ7L b5il,1PԶ2>8p w_}H 38@t"CR !]Zj@?*[7KdڒE3#i%q;gMgy"M.*,.!A9 "$+[M hIRߠze B@rgoS~LC!tO8ޝ!HMUK#v bu?JջݦV"c̕TꞞiK4!Gzd  y diꩂ~K]>\ ȃ^΀;J ل׫hMO!_O:{Tm"PpӼCEN.[;:xLf]iQ [WFdə4аR"M;^m[Y2.v3?y_[zʂN~k<'dn2,?~ H7];pdοsvY"έz_dVJ|n4y?rAj75Qkk:dw6Ua+tl'~v,vSBjw_5 &6 m[8/tO &"(.W,&z77{_ӈd!'J1.¦b֐tL! 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Tm <616C7320> Tj 1 0 0 1 352.08 615.36 Tm <53746F63687320> Tj 1 0 0 1 393.12 616.08 Tm <28537469657220> Tj 1 0 0 1 430.08 614.16 Tm <2B20> Tj 1 0 0 1 443.76 613.68 Tm <4F63687329> Tj 0.96 Tc 1 0 0 1 104.88 601.68 Tm <62657A656963686E6574656E20> Tj 1 0 0 1 186.48 600.00 Tm <4D61737462756C6C656E20> Tj 1 0 0 1 255.84 598.80 Tm <736F6C6C656E20> Tj 1 0 0 1 298.56 597.84 Tm <7369636820> Tj 1 0 0 1 331.68 597.36 Tm <647572636820> Tj 1 0 0 1 373.44 596.64 Tm <666F6C67656E646520> Tj 1 0 0 1 430.32 595.92 Tm <566F727465696C65> Tj 0.96 Tc 1 0 0 1 104.40 584.16 Tm <676567656EFC62657220> Tj 1 0 0 1 161.04 582.24 Tm <756E6D616E6970756C69657274656E20> Tj 1 0 0 1 249.60 581.04 Tm <537469656E656E20> Tj 1 0 0 1 290.64 580.32 Tm <6175737A656963686E656E3A> Tj 1.20 Tc 1 0 0 1 176.16 546.00 Tm <486572616267657365747A74657220> Tj 1 0 0 1 260.88 544.56 Tm <47657363686C6563687473746E696562> Tj 0.96 Tc 1 0 0 1 175.92 527.76 Tm <48F66865726520> Tj 1 0 0 1 216.96 527.28 Tm <5A757761636873726174656E> Tj 0.96 Tc 1 0 0 1 175.44 509.52 Tm <566572626573736572746520> Tj 1 0 0 1 240.72 508.32 Tm <46757420> Tj 1 0 0 1 259.20 507.84 Tm <74657276657277657274756E67> Tj 0.96 Tc 1 0 0 1 102.24 474.00 Tm <496E20> Tj 1 0 0 1 115.44 473.76 Tm <64657220> Tj 1 0 0 1 136.56 473.28 Tm <5363687765697A20> Tj 1 0 0 1 183.12 472.56 Tm <77756E646520> Tj 1 0 0 1 218.88 471.60 Tm <6C39374220> Tj 1 0 0 1 247.20 471.36 Tm <65696E20> Tj 1 0 0 1 267.36 470.88 Tm <766572676C65696368656E64657220> Tj 1 0 0 1 348.48 469.44 Tm <5665727375636820> Tj 1 0 0 1 393.84 468.48 Tm <7A7769736568656E20> Tj 1 0 0 1 444.72 467.52 Tm <4D756E69> Tj 0.96 Tc 1 0 0 1 102.00 455.52 Tm <756E6420> Tj 1 0 0 1 124.32 454.80 Tm <4D75636873656E20> Tj 1 0 0 1 173.04 454.08 Tm <756E74657220> Tj 1 0 0 1 204.48 453.36 Tm <64656E20> Tj 1 0 0 1 226.32 452.88 Tm <646F72746967656E20> Tj 1 0 0 1 273.36 452.16 Tm <48616C74756E67732D20> Tj 1 0 0 1 327.60 450.96 Tm <756E6420> Tj 1 0 0 1 349.20 450.72 Tm <46FC74746572756E6773626564696E67756E67656E> Tj 0.72 Tc 1 0 0 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<64657220> Tj 1 0 0 1 231.60 90.24 Tm <4D75636873656E20> Tj 1 0 0 1 281.52 89.28 Tm <616E20> Tj 1 0 0 1 299.04 89.04 Tm <64696520> Tj 1 0 0 1 320.16 88.80 Tm <737461726B20> Tj 1 0 0 1 351.84 88.32 Tm <73636877616E6B656E64656E20> Tj 1 0 0 1 430.80 87.12 Tm <7765727465> Tj 1.20 Tc 1 0 0 1 94.08 74.40 Tm <64657220> Tj 1 0 0 1 114.48 74.16 Tm <4D756E697320> Tj 1 0 0 1 148.08 73.44 Tm <73746174742E> Tj ET endstream endobj 79 0 obj 10979 endobj 80 0 obj << /Length 81 0 R >> stream -6 Mas[versuche mit "short scrotum bulls" das Intenesse einigerschweizer und Bundesdeutscher Mster enregt und sie zurNachahmung dieses Eingrif fs bewogen. Die in der schweiz alsMuchsen (Muni + Ochsen) und bei uns als Stochs (Stier + Ochs)bezeichneten Mastbullen sollen sich durch folgende Vorteilegegenber unmanipulierten Stienen auszeichnen: Herabgesetzter GeschlechtstniebHhere ZuwachsratenVerbesserte Fut terverwertung In der Schweiz wunde l97B ein vergleichender Versuch zwisehen Muniund Muchsen unter den dortigen Haltungs- und Ftterungsbedingungenmit Braunviehstieren durchgefhrt (20). Unt'ersucht wurden dieAuswirkungen des knstlichen Kryptorchismus auf die Mast- undschlachtleistung sowie auf die endokrine Hodenfunktion derJungbullen: whrend der Aufzucht bis zum t. Abschnitt derMastperiode bei einem Alter von B Monaten ergab sich eine leicht,eberlegenheit der Munis, im z. Abschnitt waren die tglichenZunahmen der Muchsen signifikant grer (taU. Z;, woraus insgesamteine hhere Zuwachsrate und eine verbesserte Futterverwertung derMuchsen resultierte.Beim Vergleich der schlachtkrper ergaben sich keine nennenswertenUnterschiede. zur Unteisuchung der Gonadenfunkrion wurden imAlter von 14 - 36 wochen l4-tgig B-stunden-Testosteron-profileerstellt. Beim Vergleich der Verlaufskurven (Abb. l) zeigt sich ein umca' zwei Monate verzgertes deutriches Einsetzen derTestosteronsekretion der Muchsenl deren werte steigen im Alter von28 wochen sprunghaft an und zeichnen sich in dieser phase durchgeringere Hormonschwankungen und signifikant hhere Minimalwerteaus. Im Alter von 32 bis l5 woehen findet eine Angreichung derTestosteronsekretion der Muchsen an die stark schwankenden werteder Munis statt. endstream endobj 81 0 obj 1719 endobj 82 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 83 0 obj << /Length 45 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj77 Do Q endstream endobj 84 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj77 77 0 R >> /Subtype 80 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 82 0 R 83 0 R 78 0 R ] >> endobj 85 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 23372 >> stream j!Cܐj[JD䶂p& G@3D^! / ', 4̲/IW_ ;@2:;.tB? m=Ǧ4jnڴK_Aߪ S}/QqI9"FV_i?M]rD8ꟿ}BO*^Zz!"Q5XC9SΦu 6hZߠ< l8nM۔Li  6444 )%THSJKNn׻Niצ;ӿXLpC$0 A~x!4#۔9x2MHWhBhGNoD|$MӯյDnWzh0& {4`@Q !7nNRiJA|̜y?o/'< oy.hQ9MےƟjh4!HC䀧A?_{tt=7np6Xdz#D]v`@t~ZWOMtۣBXoNG  \v5{[ zꮞAxNr~FA䱷`ݢ(Nn뭽vW*jp!BtO0A H HSP?q_c/V~[ڪoH?jItPd-?c~{.ԅ/oT*T/n2\dƮDj=A _K_vp${A-UURI*!Yrn܌d%Ck^i$Aǂl,6u]W+M54I$_e` Ao޿!~%dj{I$nOA [{}=9nߞ{^'$~ORUJ}%IW޸jX0 6van7kVɺ/O{(U$$4ب1 cA)6ҎB?b ZiVMm[ ޭ˰yӿM5ҵM1Nlckaڿ`m7MD!'a=S!&i&LV11` +z4ЈB,J2P&4a4kۦ%Mx !iM."}vk`%؈`CA$#AL,4;bmDDDD2A ]DD'dHADDDA*#Tj36ԇ9~JnY%72;3TD VM2pdg UiD! 6^@ӧO^ÄEwAc;Re+ uM|O_0&zh54D>}o蛻 N!SNaR `"GY^Y_'!/]y ->~?)o ŧW_x?_<]螽%O{Hҽ*I.۬o kޖn6*VPY r|4[c)!Ҡ7A0 nMlMI `H/b""6 T*aA%]qɺpCc4d_%8L$ A<0?#4G|!Yfa΄NDh9"\CH@i0i-4 A ABL`a3 ,!!saD1B$=d4x9B<fx#Փ3M*__6O4=0 }ـil0<0 ?I| )8AON4wzihL`a0  !x?:Φ".Aݦkva;T|0Ј gL:b,"ΤuBNpDO9FcHci檚iv[DDDDDDDDDDDDDDDDDA2GHF45M;BtbuTXJ?Rh:X0-r2d)L22dEX6 hN~$M eZ ȢW;Q~rMӷpx;geYɿk>M DNLt l칟3Y& j0O0s fI)/ EAu2yU4D#r7փy0OQ"^FW iB`H% Yޞ eyO#?'v26 'ovlwW"zMm]t;xV,VC@?п~Lo>XbEA8߾ "|t-)'%0nT 吮m)5d1(Ok URP!o骭A +_F'({u*am2BUt-ݥkݯ& m,l)Hz0if&;3\'5! $;a8/MWeDQ@0v`V شLY8R0#]cv65J'v*!&aA 0~i!:iil& ?Q  d`H<"""#" Xa 0B dK "#(33" -sLdϣS$I)6TFJ2B"ӜC:AG! rmg0BԂ}OĶ欤6J C94 E !`P3NA#%#A"uIhA  \wd;M;pP`%'NĘ]S [ m mqv':'O<O8Am}o~k|>Ӣ8os%bF$@ېodq 5DXm=[__Dp܏/#;_KY)8B۷o6HAܔKm{_ &KԑIt .Tfi0}}6]VApA"m $ ^E~l0K0*݈i0u]5A 2 ꚏal-ϳT 2%h0V ,pz0F sX_ɷ}kOɷ/ or ɍyVa /j%d LDB 0L&L Q(zeTʻMiT%V5AdEzaԒʃ*A"jqk%t֝AB :a6 @ ْH PeSV KZoLM7aA ʚ\M9H M)/c]7ujvH([@/T>ODID/*]&/Ii_H4nqwi߄'c]-]Jtu׬ :%]^'V~_ͿKB}/J"Kk4q[\J_zվIA/Ӆm)gG']zmoXZ^uD샮/o薟q޾K}]X0WӶaצO_x?}?I-wty&WWV_D&zD Fu[ ޖ$ՠO_I6֪V]6iwKM#OjiuA?VkIմj+i8VK{`4צVVM'VD N52" STP i1T4=d MP0`_ *bڏBJaWSLV0!e [WmWM \$ aUPAZmjji`2% jAk((aSJ&L.hIahTT0`;$`PB#B* k !gQ8 "*ЍVk-VתZ;jOJkiЭPR,WWg# e~ ={ڪ*f]=82 d"-䘤)10DE1 2 " BdTr.ʚ eaa4efM 2h3>AKMS*b."DSO !&ˤ v qV &c tMP 5 W\ SE=1iXk]5SU}V&BFG+!5U@0 „a aa5`ei:' Y0,b b,Yj#B;1P)ЋBTh0ZLjB#u_ү8؝}5`?h|- W>yz0޶C_m[&Ⱦ)C%AI$5_Zo_ֺ}- ᱵa8eCk_u6 H|a_ _&"g(ʨ9\Ly6GN,NAN;+`'!H$[Z`VMRze$ɲL.eTS-DQXFA'oP`S"q:eS@U!'B 9jui,W&ZpTM:ߑ3ՂmJ[MV\^n%HFoD Yn\^aoߝ7oiBq2@fT:3s}+z~QI_V!丄`0#A!ufEֿ=}wNhRiP{%zy/|H5]!Sv#Ƿ!!X"+nӮ_a,8dJaQ/a7H/Z _n-U]:Nm( MZʛLP//괞A'( E_d\6{*<[O+߫zl?.{WviIdX_{;!Ҝ ^ߵ _ִKޮwf{M_A[3?5=o]={x7/ AH=*\+_~z_]v]kmU5mInHZn[i/޻XK׵TW v_NV]{ Zփ\Ҹie>%ګ %l%5T8j`%צGb /414ZkL{XN@"@iSx(vD %Q[Tz`Q> c! ~I $-BMSH.Ӧ .Zh0jNWMza5L/IIp. !a(0Qib3&0LtUX2ReYEI3ZDGde`hFq"*zUKe[Uqv׈#'ν^bvP/}d&$^AV"0OMPa<&AlM:T%3*;dD]el +'}S!aB'o'4N]4xA };6AvnzveN YgtS$Nީ:eJ Ub"jl߿#KôEz'7Mk+wX |&l9vvx=ޓޓxO|gPL W:(&B#=OII?_t=i}z_7]X T8$l/Jxv_wj4 ÇvG4'!DKڢ#~o҆ 88I;]'ECC0dp6zL !GTXu ja/oa?#W{tPo0䷫}! q jݯR1נM~]m-;ۿ%_]]IWu֗6ׯY3%I/uڦI{ҶZI]iiixjk^u֛WmuUizZXPl% l&fgL4iD %JvhJdp"Bat,TCbCAx_Al NUb)b>`R"cؤza8:\iքU5UT[]H' k]Xi &xAa c GdHB׸i   d.RV x&Q`DDDZ 00`Gq#=DRD|F!T ު K ZOk 'v@Ҥ8hqx 2ja\D^a#IazʂQkge4q;z_ F#f?!o}:F aA"!ex&)r2; &l8@D4 !B M96XGZ 4 ٔ¦DI42"% tO%KhqN+KܜAh ' rnr,)r/A7C2&[.Bi7j\mTENTUz''G۶:NI߯&9!`5FA&VR zA]-$공 [* zbZAzD1"T}=z?U8ӵ ?zZiU<75dW]%}&/T!0vohQ?7($PU vIDŽ6 kUj|C/oO_LtIhg5Odr#a5ɀYI-6@y ௶}$Jm/7p=|sudGP{"=/~Gz'vI/T޵Ka~8lIK4T[ *בYIߒgk܇k8}u Fz^k$6]xe7$]붖zjVN y!z`[]m $3@]l ႮIV0a* 'iW)NA::k UD\~-ج>>fNt!V ޴QuIuMWAG,V2-L2$&ci]Ƚa+AP馼L(48D!{ ai&XaPɁX0 """" c"DhG5P_W]Mm5qG:/#u+3B-XW kUڕOI+S:ؔ:53c!Ie4-A:ރA0:Q/r>r1FNwI2k#Mפ> ujRڿK_?ri>L&[]5'd4ۿ Tӳchkcbi A`^z^L%M5VA6Iom3\~ɶU"FC2 ?Eo)TD! 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<68656C66656E2E> Tj ET endstream endobj 103 0 obj 11399 endobj 104 0 obj << /Length 105 0 R >> stream -9 Die histologische Untersuchung der Muchsenhoden zeigte einehochgradige Atrophie des Keimepithels, das Hodengewicht betrugrund ein Drittel des der Munihoden. In den von 14 Monate altenMuchsen gewonnenen Ejakula[en wurden keinerlei Spermien gefunden.Das beobachtete Deckverhalten der Muchsen unterschied sich nichtvon dem der Munis. Diese qualitative Aussage hat jedoch wenigGewicht, da keine langfrisLigen ethologischen Erhebungendurchgefhrt wurdenZiel der vorliegenden Arbeit ist esr die Gewichtsentwicklung derStochs im Vergleich zu Stieren unter den hiesigen Haltungs- undFtterungsbedingungen zu untersuchen und ber parallel dazudurchgefhrten quantifizierbaren ethologischen BeobachtungenRckschlsse auf die unterschiedlichen Gewichtszunahmen zu ziehen.Die Frage ist, in welcher Weise der knstlich herbeigefhrteKryptorchismus die Gewichtsentwicklung beeinflut. Aus den bishervorliegenden Befunden ergeben sich fr die hheren Zunahmen derStochs zwei unterschiedliche Erklrungsanstze:A. Mit dem Eingriff wird die Androgenwirkung reduziert und damitdie soziale Aktivilt, v.a. das Sexual- aben auch dasAggressionsverhalten herabgesetzt. Verminderte Unruhebedeutet geringere Energieverluste und damit verbesserteGewichtszunahmen.B. Durch den verspteten Eintritt der Pubertt, den darauf hin abersprunghaften und erhhten Anstieg der Testosteronseknetionfindet eine verst,fkte anabole Wirkung im Organismus st,at[.Sowohl eine verminderte soziale Aktivitt (siehe A.) als auch eineverzgerte Pubertt mit anschlieend verstrkter Androgenwirkung(siehe B.) mte sich, so die Annahme, in einer erkennbar verndertenAktivittsfrequenz uern. 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LiteraturEndokninologie des 5tieresHodenfunktion und endokrine Regulation Der knstlich herbeigefhrte Kryptorchismus ha[ seine unmittelbareund direkte wirkung auf den Hoden. Die im scrotum paarigangelegten Hoden erfllen eine doppelte Funktion:A. In den Samenkanlchen (Tubuli) luft die Bildung der Keimzellen'die Spermatogenese ab.B. Zwischen den Tubuli, in den Leydigschen Zwischenzellen findetdie Produktion der Hodenhormone, die Steroidhormonsynthesestatt.Abbildung 2 zeig1, wie die Hodenfunktion ber Gonadotropine derHypophyse gesteuert wird. Diese geschlechtsunspezifischen Hormonewirken eng mit den testikulren Hormonen zusammen. Die'wirkungdes follikelstimulierenden Hormons FSH ist auf die Sertolizellen dersamenkanlchen gerichtet und regt die spermotogenese an. Derwirkungsort des Luteinisierungshormons LH sind die LeydigschenZwischenzellen; es stimuliert sowohl deren Ausbildung als auch dieProduktion und Freisetzung von Tesbosberon'Die zentrale stellung in der Regulation der Sexualfunktion nimmt dasHypobhalamus.Hypophysen-Systemein.DerHypothalamusisteinBereich des Zwischenhirns, der mit dem Gonadotropin-Releasinghormon Gn-RH auf die Hirnanhangdrse einwirkt und denHypophysenvorderlappenHVLveranlat,LHundFSHabzugeben'DerHypothalamus ist einerseits die Schaltstelle, an der Umweltreize undpsychische Einf lsse verarbeitet und in hormonale signaleumgewandelt werden, andererseits stellt er das Regelzentrum dartber welches die testikulre Hormonsekretion kontrolliert wird'Dieser neurohormonale Regulationsmechanismus wird als negativerFeedback bezeichnet. Die Gonadotropine wirken ber die durch sieinduzierte Androgenproduktion auf ihne eigene Sekretion blockierendoder stimulierend ein. endstream endobj 113 0 obj 1750 endobj 114 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 115 0 obj << /Length 46 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj109 Do Q endstream endobj 116 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj109 109 0 R >> /Subtype 112 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 114 0 R 115 0 R 110 0 R ] >> endobj 117 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 27858 >> stream MAΪ?Δ,HAGZPy߯k[ z~?e9P% ? 8]n!QY[$~ZVZw.YF[T _tӓja=58mPZQ, D~MCѫ޿E4HcC=&߿ڏkdD9_CjWSBy@$8,MQ/W;4AaOӧ_,&>'Osͷ:KHƩ/צECn/%]~z|'j]2~OEW?~A_D}vUߩ9EZwZJm{ .A?{HzIsv7l/װiO_>~?'i~Jl/8`#?߶?i#,7ݯ{kZl_~$[ k޶mawN8y_US!!4A!SN&iDGTB"#״_O KX5Aܘ"""#~i]uV;bݤ !v#M-m&60V =E L@FuxzaSB!] BЈ4UM4iGɕ4-2O:bfKp% UaLdT1q兣!|.)@aM>|!ymdfu#0f`<x9sp0 i-4\C0C>x9sijwvt2p`σijoR]e,Fu"M<fx9CჀ a5Tih4 3aɹh!@?O AЅd'| ABB'N 9sa馘PL.^ӻNM4i3`(tCD&"""""#M4Ӵ ݦp }M #5""""""""#_?馚iڦL2nDDDmSOOt(}m0hNuDDnXF8Aq qP 4M4t"t `DDDDDb""""""""""""#'8c% 2h'φ)hl e4xp[9L,Ym#). \3#|phFZF9x-ٗ‘ DDDDDG尦\q"d7YM 6d]OH."N7s 5b8 as.GЯ&H0#2_6 bѰP|">=gv}2:=#?"# 3#8αLH#Q˃sG "$DDxdpm08)G_3DjDDHB ÐPjY5 dLG׿ג ub-֑$Gn!e~Y ,a 9[ WsYhV?V@n4f#o_,Kק-~ḙW_S"(}upcX[͌ș,Fǘ,CDQB]ݧ_aBFE1.?t u}o^5NWu_y7R_NiZkb2| i!uc":0E ?iGdqs?$U5mzi0B,vDq`7 T$lO2#?"#elpDDDC 'rAo@Ag(39?-YLR 2YliW{_E '0Mp__Po~vXD5 ^jL T&A|I؄A4!$h=^_<'xO/iq)rVLBEbN%ddaeZ_"<"ݠ%̈́/ G & Ӡ tEwzގnGa8R=pM:Gh6Bub~D~dw!i?`:DG o[WoĖn[ٰu_ la}(di_(0?uq-0}l ݯ~a(0/k_T'Td+A,/m+6 eyn[NaJ+ )(8T-2Y %a0D8vVPdW1KqA05%^a#KȘb#`:#_jEqAF`K DHsb?Ȯw!9@Aȇ(س.׿] 3CO!K_O]_Z v* }ޙPˍ^w) (q*Me0A}SMc)Ĉ!ݢ~@@LU@&@i%N|*NM?DȹImXx/TOh M ˚A7fbmD=f9:z{!GM.ܛ,Dy]id - &`\H"˂ (C¸`#X_2`- :@b`3Y㑎C,c3la;k089'/`ak_D2C.H!3Zn_vBq&yr׈@M)_ٍ5_2 OߵKNk\~_g|?vׇ i_r2GTj Π`.!(AߢP4rDX$@)0PgNH pCeD#A &!uhD3Gϋ;q_Ѓ_&w'`%D[5"jm9k؃o; k_omΎMEm.,˚&C&vBbޛ! 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Rege Iz entr en Releasing- GonadotroPe-Hormone Hormone Ge-chts-eIn SexuaI-ho rmo ne Er fo lgs-organe ll'lTnhibi It Sertoli-l sfermato-zellen I geneseIIABP), I l::l i.tt"'Androgenel-l organ(strogene) luus*e Gn-RH HVL FSH ,/Leydigzellen I ( aus KRAUSSLTCH, 198 1 ) Fr FSH, das zunchsl durch das gemeinsame Releasinghormonpanallel zu LH ausgeschttet wird, besteht ein weit,erer Regelkreiszur Feinsteuerung: Die durch FSH beeinfluten Sertolizellen bildenInhibin, das die Abgabe von FSH durch die Hypohpyse hemmt. :Auendem produzieren die Serlolizellen unter Einwirkung von FSHdas androgenbindende Pnotein ABP. Dieses Protein schtztTeslosleron vor einer Verstoffwechseltrng und transportiert es in dieTubuli, wo es anregend auf die Spermorrtogenese wirkt. Versuche mithypophysektomierten Tieren haben geatrigt, da durch Verabreiehungvon Testosteron die Spermatogenese attfrecht erhalten werden kann.Das lt darauf schlieen, da Testostnron selbst den entscheidendenKontrollfakton der Spermatogenoso darstellt. endstream endobj 121 0 obj 1115 endobj 122 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 123 0 obj << /Length 46 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj117 Do Q endstream endobj 124 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj117 117 0 R >> /Subtype 120 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 122 0 R 123 0 R 118 0 R ] >> endobj 125 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 40850 >> stream l,7j~Ptaaϔ _??Sz.? ~OөmBw U!h?YkZL'YyAj?/bEK$I [T!_iheG1S ~5뽝*?MAX%xj|'p@X)5I?(,%J (MZ^󾠄HdXgeړ+2di ՄM;i9}<i%|7_ރ{[󰖱AjY #$g!#0&A_(g"B+Mi=k!#P<C?uG@! 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Beeinflussung desendokrinen Regelkreises Im Sinn der Rckkopplung ist das Hypothalamus-Hypophysen-Systemfunktionell von den Androgenen der Leydigzellen abhngig. GeringeDosen von exogen zugefhrlen Androgenen unterdrcken diehypophysale Gonadotropinausschttung und haben so indirekt eineAtrophie der Leydigzellen, eine Beeintrchtigung der Spermatogenesesowie einen Gewichtsverlust der Hoden und anderen Sekundrorganezur Folge. Hohe Dosen exogener Androgene bewirken zwar ebenfallseine Hemmung der Hypohphysenf unktion, sie stimulieren jedochdirekt die Tubuli und Sekundrorgane (12).Man konnte feststellen, da nach Abbruch einer lngerenTestosteronbehandlung die primre Hemmung der endokrinenHodenfunktion nicht nur aufgehoben wunde, sondern in eineHyperfunktion umschlug. Diesen Vorgang bezeichnet man alsRebound-Effekt. Man fhrt ihn auf eine Desensibilisierung desHypophysen-Zwischenhirn-Systems gegenber der ursprnglichenHemmdosis der Sexagene bei langfristiger Zufhrung zurck (22). 3.I.J. Folgen des Kryptorchismus Das augenflligste Merkmal des Kryptorchismus is[ die geringereGre und die Schlaf fheit der dystopen Hoden verglichen mitNormalhoden. Die deutliche Grenabnahme wird auf dasSchrumpfen der Tbuli zurckgefhrt (KARG, f950). Folge diesesDegenerationsprozesses ist eine Asperime. Der knstlich induzienteKryptorchismus kann deshalb auch als Sterilisation bezeichnetwerden. 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0 1 206.16 54.00 Tm <7769726420> Tj 1 0 0 1 232.56 54.00 Tm <64617320> Tj 1 0 0 1 253.20 54.24 Tm <47657363686C6563687420> Tj 1 0 0 1 314.88 54.48 Tm <6665736C67656C67742E20> Tj 1 0 0 1 368.40 54.24 Tm <44696573657320> Tj 1 0 0 1 405.60 54.72 Tm <7769726420> Tj 1 0 0 1 432.00 54.48 Tm <616C73> Tj ET endstream endobj 135 0 obj 12750 endobj 136 0 obj << /Length 137 0 R >> stream T I II II T TI t t I I I t I I I I I I I I - 13 c/eF Minderung der FleischqualitL durch Geschlechtsgeruch vonpraktischer Konsequenz. GLOVER (f956) berichtet von Kaninchen,bei denen durch scrotale Hitzeeinwirkung die Fruktose-Konzentrationim Samen anstieg; diese gilt als IndikaLor fr die Androgenproduktion,was fr eine Stimulierung.der Leydigzellen durch erhhte Temperatursprechen wrde (?2). Andere Befunde sprechen jedoch fr eineEinschrnkung den Androgenproduktion beim Kryptorchismus. KARG(I96t) fand in einem Versuch mit Ratten signifikant ' niedrigereVesikulardrsengewichte der kryptonchiden Tiere. Dieseakzessorische Sexualdrse wird als der klassische Indikator derAndrogenleistung angesehen. Der Differenzienungsgrad derLeydigzellen und damit die androgene Seknetionskapazitt war beiden Abdominalhoden regressiv beeinflut. Die signifikant erhhtenLH-Gehalte in den Hypophysen der behandelten Tiere zeigen, daO essich beim Abfall der Androgensekretion um eine vermiMerte LH-Wirkung am Rezeptor handelt (f l). LEUCH (1979) folgert aus seinenUntensuchungen an kryptorchiden Stieren, da sich dieEmpfindlichkeit des Zwischenhirns gegenber der regulierendenWirkung von Testosteron verndent. Er vertritt die Auffassung, dazur Ausbildung eines funktionstchtigen Feed-Back-Mechanismus eintestikulrer Faktor whrend einen bestimmten kritischen Zeitspanneauf den Hypothalamus einwirken mu; der durch die Manipulationausgelste temporre Kastrationseffekt hat zur Folge, da sich derSteroid-Rckkoppelungsmechanismus nur ungengend entwickelnkann. Dies fhnt einerseits zu einer Erhhung des Schwellenwertes frSteroidhormone am Hypothalamus und anderenseits zu einer erhhtenEmpfindlichkeit der Leydigzellen fr LH whrend der Pubertt unddamit zu einer erhhten Testosteronwirkung (20). t.I.4. EntwicklungderGeschlechtsfunktionen 1.1.4.I. Prnatale 5exualenLwicklung Bereits bei Vereinigung der mnnlichen Samenzelle mit derweiblichen Eizelle wird das Geschlecht feslgelgt. 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Handelt es sich um einengenetisch mnnlichen Embryo, so wird beim Rind schon ab dem 40.Tag das Gonadengeschlecht festgelegt, indem sich die Hodenstrukturzu differenzieren beginnt und Int,ens[itialzellen angelegt werden, diedas Hodenhormon Androgen produzieren. Dunch Anwesenheit dieserf talen Androgene dif f enenzieren sich die sekundrenGeschlechtsorgane, das somaLische Geschlecht, heraus. Im gleichenSinne wird auch das hypothalamische Geschlecht geprgt und daspsychische Geschlecht festgelegt.Diese wichtigen, durch Sexualhormone gesteuerten Entwicklungs-schritte, die Diff erenzierung der Fortpf lanzungsorgane und desSexualzentrums, finden also bereits im prnatalen Stadium statt.t.I.4.2. Postnatale SexualentwicklungDeutlich wirksam wird diese gesehlechtsspezifische Differenzierungjedoch erst im Laufe der postnatalen Entwicklung. Die Pubertt wirdals die Periode in der Entwicklung eines Individuums definiert, in denes die Fhigkeit zur Fortpflanzung erlangt. In einem 1969 erstelltenGutachten ber den Eintnitt den Geschlechtsreife bei Jungbullen wirdder Beginn der Samenbildung zwischen dem 5. und 7. Lebensmonatangegeben, die Produktion von mnnlichen Keimdrsenhormonensetzt jedoch bereits frher ein (7). Charakteristisch fr die Puberttist die venmehrte Sekretion von Sexualhormonen, wodurch die volleAusbildung der sekundren Geschlechtsorgane und -merkmaleermglicht wird. Durch das Zusammenwirken von Testosteron mitFSH setzt die Spermatogenese ein. Auslsend f r denPuberttsvofgang sind Verndenungen sowohl im ZentralnervensystemZNS als auch in der Hypophyse und in den Gonaden. KARG definiertdie Pubertt als einen Vorgang der Desensibilisierung derRegelzentren, wodurch die am Regelkneis beteiligten Hormone aufein hheres Homostaseniveau angehoben werden (15). 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Dies lt darauf schlieen, da bereils im Verlaufder prpuberalen Entwicklung eine Desensibilisierung desHypothalamus gegenber der gonadotropinhemmenden Wirkung desTestostenon eintritt (5). Eine fortgesetzLe Stimulierung durchGonadotropine induziert die Reifung der Leydigzellen, so da dieTestosteronproduktion stark ansteigt. Durch eine enhhteRezeptorkapazitt fr Gonadotropine wird deren Wirkung an denGonaden erhht. Die Vorgnqe der Pubertt sind in Abbildung 3schematiseh dargeste llt. Abbildung 3:Vernderungen im endokrinen Regelsystem whrend der Pubentt I T I I t I I II Gonadensteroide niedriq - unverndert- Erwachsenen-niv e au sensibet-abnehmende -desens ibi f i-Sensibilitt siertniedrig -zunehmend -Erwachsenen- n ]-veau Iamyse Gon aden-s ekrete FSHe Fe e db ackGo nadot rop i ne Hvp Hy us othalamIpophyselll FSHuon ade us tha Hvp Hy {,ophl,tttllVnad-o- vo vor derPubertt Beginn derP ube rt t Gonade (aus KNrG, 1974) Erwachsen endstream endobj 153 0 obj 1183 endobj 154 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 155 0 obj << /Length 46 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj149 Do Q endstream endobj 156 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj149 149 0 R >> /Subtype 152 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 154 0 R 155 0 R 150 0 R ] >> endobj 157 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 37938 >> stream *+"#8s7# 2DDDD츇\`8 .q`8~RRLXɃiArj  .eԌ$?Ɏ-W8btʀ%TMaPUKԵZ w R4&up;X|~m҄VDDGDC)޷Γ$AGIМn?`GY}r>o'68h4G;&%Ks^~/X\zHK%Ou"EvݢP=>G ]Nq pBi2cnOAOAOMu'mAAzzzz xA| mnA ɆN%= FNoݾ}I[I뤛&}I? 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T It I I I - 16 Neben ihrer entscheidenden Rolle bei der Geschlechtsdifferenzierung,ihrer direkten Wirkung auf die primren und sekundnen Geschlechts-orQane7 liegt die Aufgabe der Androgene in der Ausbildung weiterersekundrer Geschlechtsmerkmale. Psychische Sexualfunktionen, wiedie Libido und weitergehend das Sexualverhalten knnen sich erstunter der konditionierenden Wirkung der Androgene manifestieren(t B). I I I I I I I IT 3.2.t.2.1. Verhalten des StieresGrundlagen der Ethologie Das Sexualverhalten ist einer von vielen Funktionsbereichen aus demVerhaltensrepertoire des Stieres. Diese Summe der verschiedenenVerhaltensweisen lt sich nach unterschiedlichen.biologischenFunktionen in Funktionsbereiche untergliedern, z.B. in Ausruh-, Fre-oder Fortpflanzungsverhalten. Jeder Funktionsbereich fatVerhaltensweisen gleicher Funktion zusammen; er hat seine eigenenSchlsselreize und fhrt zu einer bestimmten triebverzehrendenEndhandlung (2t). Diese funktionsbezogene Einteilung erklrt alleinnoch nicht sehr viel. Weil einige Verhaltungsweisen in mehrerenFunktionsbereichen erscheinen, z. B. Stoen sowohl beim Kampf- alsaueh beim Sexualverhalten, ist es notwendig, einen Aktionskatalog,das Ethogramm r zu erstellen: Diese Bestandsaufnahme allerauftnetenden Verhaltensweisen beinhaltet deren Aufzhlung undDef inition. Zur Beschreibung gehrt auch die raumzeitlicheAufeinanderfolge bestimmter Verhaltensformen (23). DurchBencksichtigung der Reihenfolge und Erstellung vonVerhaitenssequenzen ist eine bestimmte Verhaltensweise besser zubeurteilen und einzuordnen. Die berschneidung einzelnerFunktionskreise ergibt sich jedoch auch wegen gleicher Ursachen: Sohaben z. B. Hormone eine multiple Wirkung. 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Es ist Anspruch und Aufgabe derEthologie, sich mit allen Aspekten des Verhallensr mil seinenqualitativen, quant,itativen, raum- und zeitbezoqenen Eigenschaftenzu beschftigen und die verschiedenen endogenen und exogenenFaktoren zu bercksichtigen. Verhalten kann als sichtbare undkomplexe'Aktivitt definiert werden. Diese erkennbaren Reaktionendes Tieres knnen sowohl die Folge eines inneren Zustandes sein, alsauch durch uere Reize hervorgerufen werden. Meist sind beideKomponenten beteiligt. Der Reiz wirkt ber das ZNS direkt aufbestimmte Muskeln oder/und auf endokrine Drsen. Diese zweigrundstzlich mglichen Steuerungsmechanismen zur Ordnung undKoordinierung von Verhaltenselementen unterscheiden sich v.a. inihrer Wirkungsgeschwindigkeit und Wirkungsdauer: Das Nervensystemarbeitet sehr schnell und ist fr kurzfristige Reaktionsvorgngezustndig; Hormone wirken dagegen vergleichsweise langsamr knnen aber einen lnger anhaltenden Einflu ausben (I0). Beide Steuerungssysteme zeigen, da dem Verhalten physiologische undmorphologische Gegebenheiten zu Grunde liegen. SCHEIBEbezeichnet sie als die anatomisch-funktionellen Grundlagen desZustandsverhaltens. Die Motivationen resultieren dabei aus innerenUngleichgewichten, die ber ein in die Umwel[ wirkendes Verhaltenwieder eingeregelt werden (25) (Abb. 4). Dieses Verhalten lt sich indas spezifische Such- oder Appetenzverhalten und die abschlieendetriebverzehnende Endhandlung einteilen. 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Verhalten kann ebenso wie Farbeund Form als Anpassung einer Tierart an ihre Umwelt verstandenwerden. Diese Anpassung erfolgt einerseits durch genetischfestgelegte Instinkte - wenn auf einen bestimmten Reiz immer diegleiche optimale Verhaltensantwort folgen mu - in variablerUmwelt, andererseits durch individuell gesammelte Erfahrung oderdurch Tradieren von Verhalten, das andere Tiere gelernt, haben(BEILHARZ, 78) (24). Mit der Zchtung auf bestimmteLeistungsmerkmale haben sich zwischen den verschiedenenNutzungsrassen auch die Verhaltensmerkmale unterschiedlichentwickelt. So stellt SAMBRAUS verallgemeinernd fest, daMastrassen vergleichsweise phlegmatisch und grazile Rassen agil sind(23). Die Verschiedenheiten kommen of t im sogenanntenTemperament zum Ausdnuck. 3.2.2. Sozialverhalten Als Sozialverhalten wird jedes Verhalten zwischen Artgenossenbezeichnet (10), also all jenes Verhalten, das sich an Artgenossenrichtet und das als Reaktion auf Auerungen von Artgenossenerscheint. Die Form der in Gemeinschaft lebenden Rinder wind alsindividualisierler Verband bezeichnel (24). Als HerdenLien zeigt dasRind eine ganze Reihe von sozialen Kontakten: Die Tiere beriechenund belecken sich, sie betreiben soziale Hautpflege und lernen sichauf diese Weise kennen. Ebenso friedfertig unteneinander sind dieTiere bei gemeinsamer Nahrungsaufnahme und beim Ruhen, solangekeine Konkurrenzsituationen auf treten. Dieses harmonischeZusammenleben beruht auf einer sozialen Rangordnung. Jedem Tierist seine soziale Stellung zu jedem andenen Herdenmitglied bekannt,und es verhlt sich bei Begegnungen entsprechend (2t). 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KampfverhaltenSoziale Auseinandersetzungen, die sich in gerichtelen Aggressionender Artgenossen untereinandern uern, dienen der Schaffung oderErhaltung der sozialen Rangordnung. lnnerhalb der Jungbullengruppenist das gegenseitige Krftemessen ein Bedrfnis, das nach LORENZals angeborene Verhaltensweise auf die Wildform unseren Hausrinderzurckgeht. Spielverhalten, wie z.B. Kampfspiele der Klber, dientzunchst dem Einben bestimmter Verhaltensweisen. Mit Eintritt derGeschlechtsreife f indet eine gewisse Auswahl der Spiel- undKampfpartner statt. Man kann darin bereits die Anstze zurAufstellung einer Rangordnung sehen (23). Erst im Alter vonL L12 Jahren sind die Dominanzverhltnisse soweit geklrt, da zurAufrechterhaltung der Rangordnung Drohgebrden gengen. Um ihreStellung zu verbessenn, fordern die rangniederen Tiere ihneranghheren Artgenossen immer wieder heraus und verwickeln sie inKmpfe. Soziale Auseinandersetzungen finden in gehuftem Mae inKonkurrenzsituationen statt. Kurz vor, whrend und nach derFtterung steigt die Frequenz aggressiver Aktivitten s[ark an. Siewird auch durch Strungen von auen erhht. Kampfhandlungenwirken ansteckend: Die Artgenossen beteiligen sich entweder amstattfindenden Kampf, oder sie greifen andene an. Rinder kmpfen,indem sie in Kopf -an-Kopf-Stellung versuchen, den Gegnerzurckzudrngen. Diese Kraftprobe, der meist ein Imponier- oderDrohverhalten vorausgeht, wind als Hornen bezeichneL.Ziel des Schiebekampfes ist esr den zurckweichenden Gegner zuunt,erfahren und auszuhebeln. Gelingt dies, so hat der unterlegeneKontrahent keinerlei Mglichkeit der Gegenwehr; en wird durchKopfst.e gegen Schulter und Flanke in die Flucht geschlagen. DasHornen, an dem beide Tiere gleichermaen beteiligt sind, dient demgegenseitigen Krftemessen. Das Stoen ist dagegen eine einseitigaqgressive Handlung, die der DemonstnaLion der berlegenheit oderder Herausforderung zum Kampf dient. endstream endobj 193 0 obj 1953 endobj 194 0 obj << /Length 27 >> stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 195 0 obj << /Length 46 >> stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj189 Do Q endstream endobj 196 0 obj << /Type /Page /MediaBox [ 0 0 595.20 841.68 ] /Parent 11 0 R /Resources << /Font << /F3 5 0 R >> /XObject << /Obj189 189 0 R >> /Subtype 192 0 R /ProcSet [ /PDF /Text /ImageB /ImageC /ImageI ] >> /Contents [ 194 0 R 195 0 R 190 0 R ] >> endobj 197 0 obj << /Type /XObject /Subtype /Image /Width 2480 /Height 3507 /BitsPerComponent 1 /ColorSpace /DeviceGray /Filter /CCITTFaxDecode /DecodeParms << /K -1 /Columns 2480 >> /Length 33580 >> stream [&O ^5eȪ>ʯ=z r t[|u-qf"!}4މÿ@m{Zk C~?6Xh_XB_  p#}/?|aYL/=.[`WǏe/)eZ5z#@XJr?@@y_+Ƞ8S{Wߎ_*-ih?}{Kʐcz_,`HDPL w0jL&!gVHva=;7'oߪNh4?_MM{J_3K7)gb.ʱ "3pD9H#!i$ii0BDAXFU$nöIA4!m.R8m{IIYAK齵= s$æޫIt7;m6w_$/kǾq޿Jٸ+^ޫx1ZT-u}I$~Mpҥ0hG\WH:?WZIS"0'K$]=vd$o8v߄]mt%B/6m$mz_AUI,Z{kك}'LW A'54T =+jAbikkBJI DPT-aa?ׯOׯ y 9X DA;ʪ;*^ \: ,ꪚ 'v3rm5; vwIUS]V 3r})8]-=4HUMa itKR4+ QwJƿuWCH"0hH;"2 k@8NCI)(&h26$A[4D)DK?flNGG!J y!uIS@@ ah4jpdLѱ"C$@E  r AÚ@5L5&J~v<' xN!Ҵa4!ЋA'aPoP( B4a3<F"(Br2#*_i"sM Nԏ4aZ^NGcDvㇲ((ia?Af`$_I.Ԟd@$sǠAA/"[@>hdsT͢st$tKLmڧvеM?7AKi}6~H7O֭m7%ʹHw[F9ݻ%4u]]{T_Z]]oSl8NzGjo_7"uC~zKj魽zow>k㵫ص]wvKz ?p{c_hvҽAd,ay޿TֵuZxw~7oz8c_mb_z~MS__:G3 }K_Woߓ"~~ |ph<}we"GD}//?! ׯ/_D绤(+(ZWk+_(^I7{/_$e&;v]p`[iYV{am[ /kkmaIz_u?mXQLTT?cb⫺K8=t}}q_v8j_m$z~bc pӻl_~WVmiivVC \h0iL{"0NOwv ~bc *-W^8`D!UL KM4"0!-t AK%MoOt߿w_$]_z>t{w Wkc zi/ƾj?۽κ}w_~B~6/[_e"?k[V}a\vC~6~j>fۏY{UKa>ma!i_ oQ[?#n"M/~ WX9P_.,tpnXk3!?Oo ˿Rғ$_t-?):?߯J--_׽_Mwӿ!H$_??TWҴ.MeEv"_r}dt 6 .l*Wmm.ia?i{}u Jׯ~E~LTS X# -j[#J- \%^uskvVՄAʀwi}V7*ئ67دbllVaҴJv0k {mH2 xiAhZd[M AꝦ4 M+im8ب؄1[ӏ"]lRxd#.hX'*05DL4D Zjj\Ӵik"$fe- [h0Aa0A.P0& !#HD!'KTua-.9:rzIŅ.#嘰̄[,2W&dePr!Vva0XM0ALڰr\$UO4 efF%DM?ɔoi4"R#C^(i}M p]}Ziokt#b_ۿAt]VPFU]>vWOB t* !h_ "NtD>,0)!YȐ!d$Pr "+ߠN|K}0d<0@ =<.uifa r}3 0?C8a0Mt"[*eЇ h2@CQ)@ȁd0E; NӰ`Iޝ B SNF;oN;TvM4[ 0D0K y tA?\2" $#v>hQ%O}Izz}]{IwۧoKnݧwIˣH'i>{Ҿ]TOT߽}~:>i/_kQIҺ+Zwݻ>G ?w"?0G ]m}Y0_ֿ|l\/200~YQDIԿ%ο}VAZ$z GNwmNH$_޹s[W]v.ki:vMvW/omouuM!S-=&+N_VӾհ0 . 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SexualverhaltenKopfste sind auch im Zusammenhang mit dem Sexualverhalten zubeobachten: Aufspr