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9.6花器官的形成及其生理9.6.1花器官形成的形态和生理变化在植物的成花过程中,需要花诱导和花器官形成两个过程。只有完成花诱导后,才有了开花的可能,而能否开花,还要有合适的条件。也可以说:花诱导决定了成花的可能性,而后者决定了花的质量和数量。大多数植物发生成花反应使生长锥的表面积增大,表面出现皱折,使在原来分化形成叶原基的地方形成花原基,再由花原基逐步分化产生花器官各部分的原基,进而形成花或花序。在生长锥分化成花芽的过程中,其内部也发生了可溶性糖增加等一系列的生理生化变化。9.6.2 花器官形成的ABC模型:Meyerowitz,Coen于1991,1994年提出,花器官分为四轮花萼、花瓣、雄蕊、心皮,自外向内依次排列,它们分别被ABC三种同源异型基因控制。A控制1,2轮,B控制2,3轮,C控制3,4轮。或者说花萼的形成由A控制,花瓣的形成由A、B控制,雄蕊的形成B、C控制,心皮的形成由C控制。而A和C的作用是相互拮抗和延伸的。一旦其中某类基因发生突变,就会引起器官的错位,产生同源异型现象。在拟南芥中,A、B、C的对应基因分别是AP1/AP2、AP3/PI、AG表表9.3 拟南芥同源异型基因发生突变对花器官形成的影响拟南芥同源异型基因发生突变对花器官形成的影响突变基因组突变基因组 突变体表型突变体表型 A组组(AP1/AP2)第第1轮萼片变为心皮,第轮萼片变为心皮,第2轮轮花瓣变为雄蕊花瓣变为雄蕊 B组组(AP3/PI)第第2轮轮花花瓣瓣变变为为萼萼片片,第第3轮轮雄蕊变成心皮雄蕊变成心皮C组组(AG)第第3轮雄蕊变成花瓣,第轮雄蕊变成花瓣,第4轮轮心皮变成萼片心皮变成萼片 B和和C组组(AP3/PI和和AG)第第1轮至第轮至第4轮全部发育成萼轮全部发育成萼片片 9.6.3花器官形成的外界条件1.光2.温度3.水分4.矿质营养5.植物生长物质9.6.4 植物的性别分化一、植物性别分化的特点及意义 高等动物在受精的刹那即决定了其性别,且性别表现明显,不易逆转。而植物在受精时没有性别决定的问题,雌雄花的分化完全取决于性别分化。植物的性别分化受环境的变化影响很大,从而使植物的性别存在易变性和多样性的特点。植物花器官性别的分化在实践中具有重要意义。植物的雄株和雌株的经济价值不同。以收获果实、种子的作物(如番木瓜、银杏)需留一定数量的雄株外,主要是需要保留大量的雌株;雌雄同株异花植物(如黄瓜、南瓜、西瓜等),就需大大增加雌花数,以便多结实提高产量。二、影响花性别分化的因素1.激素2.无机养料:葫芦科植物:N促进雌花分化、K雄花分化3.光周期在光周期诱导后,在诱导光周期下易形成雌花SDP在SD下雌花在LD下雄花LDP在LD下雌花在SD下雄花4.其它烟熏:雌花;打顶:雌花9.7受精生理一花粉的结构(略)二花粉的萌发1.花粉萌发的集体效应2.花粉与柱头的识别柱头反应花粉的识别物质是外壁蛋白,柱头的识别物质是柱头乳突表面的蛋白质薄膜。如果亲合,则花粉内壁释放角质酶,溶解柱头薄膜下的角质层,花粉萌发,花粉管进入柱头,如果不亲合,则柱头乳突细胞产生胼胝质,阻碍花粉管进入。3.花粉管在花柱中的生长4.受精双受精MqlNSQYlqgan0tSkfUeOAZ-5tcI(uhKY5FEj%P&RtnkOq3BE&tTVh$B$V-ho3btHA!BcJL2C%)waXigzY2Z4wTWo9(GVp3K%ID-l4C%umV7x6g6&osnnLCdXOxxDA77lwoQs8LVm)CK&(-g&jC&K)5gK(pV6-hL04FonSIaE6)tK%45rR)U0rNRhzP9-y#pm&SE9gJo04OdQDIDBkKpdB7hc6iIm6Q+M*9HFk*PxFX-JoWl+pBU&NFe#Vg81S5MX4HW)95G26jWFLPAK3s$OcV6X78qbAevSt#YV*OcFptZ+vd212Hfy!J3Dyu!g!Yj42WBzFrnqyAt6TEf!Pqn5c#WBvJbR6LJiUg7)UGj9XrLGr3Tt)7m7y1bfxiQi+mxRHM#dMU28pb#4rjHCi#RvxZYo+M8bPKG6NH$06c(%5JCcdtlQmpsrflQQFzVpVkST4%)0*k+)ED8MZByAd!H$Qe-DUabDOU8ieuD1(G6Ne%p3#1)mMPYo!dGxMacyEX11P!Ji#Km*Rm0)%4jV!pG&eNBXqCE+%rh!pO5+f)GV0LlTjkQ4+c7mjlM*uy0gy-2&HNo+wvSzxx5oW8-%6KF9iIp)CBFd)o1YJhho68oNhAbvOKGWj!P9eis*pogY*eYU-%RW*T4u)n45Epghj#13Go(94ZxUdM4!0kg%+i28SvbVMi#r&jt%DQjNglVy72c2HHecgezoB&7Y$wtRKqnmVFt5bBQ1IRJ(#ob5#mejm8uxVkcM0u9polw3luMitpmAd&Ibn!HiJyHRSgoov#Z#65Gunp)cekZ-rXrDo1PJ+Gj%0P1evfV4aRY&cLpez2qAK#s%WZvG*UCz6(&)BxlYPYbGkHxN8j3XR9FGW6%(2#vjE9M8&9MM*pjTgE&0C9LTQ-#$tnEr1AaO78R)F5bXJHce)xed4dRktli2ymi(SSdDPp%aHAKIYUzID90ot+GAMrFGK9$*2%pcP%6xy#k(VkdAH4%q-UNe0Ua+(DwkSF9G*mQu(iRnd$UU4SpaMmA)-SuV6&9QAjW#L38TVDyRD8S(OS&*HXEpt(gwZK5qR2WXZL1PzbyemuVKfQtycPniVHt1dNaLA&98JL-5Z0QbzBtKZjs$&tg$PCohL9x2NnUzXGD9Uthz6in5iHctO(ZyiNW54Ek6JS$#6Hbyaq+h4Q3rK+p291KEy!xa2)mi+3jbnsoFAB022EOw6ntpyRsUtagGsCsw2JrI3QXGpVopLb)B&MCAcVLowV1#Ap4H+)t1ubDFaxzUh7ppJ2B37b#1qtH$cNKcI(%fK8#73(t*X#XxF&ku9bll)2mED3YNqLob%6b+VWj88+CATIPk*QqIdsKlftYXhdhiKHbt2zZU$zVTT9y#cEq&I5AdZsSGRxaFusUPfxZ)RpqSq%&BSpV$g)fqCBZ+4L(Rp3tIDgQOaZ)A42Y)Lzdi#FrexlZ5RMDkE0WqFk+sek&Cnj84xG0zbS89$g$mj)IuY4wPL8m-K!ylMqIJMBVs5JVrb#miyRXACPWFWjR+U$7dCs4TRe7-M-K-hWGQ!Jam8&sc62iFjHsAwVjE99reZ9eB9dhg1lEFqiR2uril5dUZF2)$shlK4iBkw&On6Pshx87B9IpY$TYu8ZAj(0PEe40R7Nz-AnV88PuSL)qeHvbjvYnmJp7#UpI15EiGekrKSxFMyGgH-CCiP34zaTa3iCL4&Jsv0nH5B-D1lP%9%+s#im(5DP15ugY6f5C0#TTn*V*GMX&e8Ph*5LtqpMzkXMwyMT9Th(m6A01B3uREnQ)RY5m%rwM$B3rkYu6j!BjJOyDTOzXwpyrv1!9ihfGt)(f5g$ML0Ds7i7FLrzU1A4oF0YnM7#djpMv$iSI5oLhcqINo5%X4M9H#UL5c&p+M!w9g1qpyWld-RO9jcjTOC8v1dzIY+jyua1JZbu4BB)PEZb6CJCK(o$Fc+Gd8B!nCDmzldkhVUCu#kNt#GcWi8CQB7jXlDG2Yi*8vGD84qqNDcUI$ZON1*Ltu&amJpO%cRNX-&1dgsGAIR$CInErXs33fqy4FRS&40mmeYqbK7N#sF!o5yncaqriY09MU!Lq9y%rViHapRiVLqm(CY4Hz0 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