马尾松枝叶特征与其对松突圆蚧抗性的关系
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摘要
植物对昆虫的侵害不是被动地忍受,而是通过许多种形质指标结构和生理过程对其做出积极的反应。由于抗虫树种具有独特的优点,因而近年来选择对目的害虫有抗性的树种受到重视。本文通过对马尾松枝条和针叶外部形质指标的测量,并应用石蜡切片法和超薄切片法分别来研究针叶的组织结构和超微结构,分析其与马尾松对松突圆蚧抗性的关系,得出如下结论:
1.未受害情况下,高抗家系的分枝数多于中抗家系和高感家系的分枝数,高感家系的主枝和侧枝均比高抗家系和中抗家系粗,由此推测主枝和侧枝越粗越容易受松突圆蚧为害。高感家系的针叶长度显著大于中抗家系和高抗家系,表明针叶长度与抗虫性呈负相关。高抗家系和中抗家系的针叶容重显著大于高感家系,表明容重越大,抗性越强。受松突圆蚧危害后,高抗家系和中抗家系的松针保有率均比高感家系的松针保有率高。受害后,除高感家系469的主枝和侧枝比未受害组细以外,其它家系受害后均比未受害时增粗。受虫害后,高抗家系330和中抗家系155的松针保有率没有减少外,其它家系的松针保有率下降,说明抗性越强,松针保有率下降值越小。
2.针叶组织构造比较结果表明,在未受松突圆蚧为害情况下,角质膜厚度、下表皮厚度、表皮层厚度和表皮及角质厚度厚度、叶肉组织厚度、树脂道数目和直径、维管束直径与抗虫性呈正相关,而上表皮厚度、气孔密度、内皮层厚度与抗虫性无关。受害植株中受害针叶与未受害针叶相比,具有抗性的家系受害针叶表现出角质膜、下表皮、表皮层和角质及上表皮显著增厚或不变化,而感虫家系表现为极显著变薄。未受害植株的针叶和受害植株中未受害针叶相比,具有抗性的家系受害植株中未受害针叶的角质膜表现出增厚,而感虫性家系角质膜变薄,受害植株中未受害针叶主要通过角质膜的增厚来阻碍松突圆蚧的入侵。
从显微结构上看,高抗家系松突圆蚧吸食处有堆积物出现,未受害处表皮结构、叶肉细胞、树脂道和内皮层等组织结构均正常;中抗家系表皮破裂处比高抗家系增多,且叶肉细胞分界不明显,而内皮层、维管束等结构正常;而高感家系上表皮细胞和叶肉细胞变形、缢缩或破裂,树脂道变形,内皮层缢缩且细胞内内含物减少。不同抗性家系受害植株中未受害针叶与未受害植株针叶的组织结构相比,表皮结构、树脂道、内皮层、维管束都未有变化,高感家系叶肉细胞中细胞器急剧减少细胞内有沉积物出现。
3.通过透射电镜拍摄的电镜图片可以看出,受松突圆蚧为害后,高抗家系受害针叶的叶绿体片层没有正常的清晰,高尔基体增多,小泡大量形成,并且有嗜锇颗粒存在,表明高抗家系进入病态。受松突圆蚧危害后,中抗家系马尾松叶绿体片层部分消失,嗜锇物质大量存在,部分叶绿体被膜破裂,表明中抗家系的受害程度比高抗家系加重。高感家系受害针叶的叶绿体片层彻底消失,较多的叶绿体被膜破裂,从而使光反应和暗反应无法正常进行。同时,很多淀粉粒撑破叶绿体被膜,游离到细胞质中,淀粉粒的膨大可能与叶绿体膜的崩溃有关。细胞质中大量电子致密小球的出现,有可能是松突圆蚧在吸食细胞液的同时,还分泌一些毒素来破坏细胞的正常代谢机制。
高抗家系受害植株中未受害针叶中,叶绿体基质片层和基粒片层分界明显,堆叠整齐,线粒体、细胞质等结构正常,但叶绿体外膜有少量的嗜锇颗粒存在;部分淀粉粒增大并运出叶绿体。中抗家系受害植株中未受害针叶中,叶绿体变形,片层部分模糊,叶绿体被膜周围有大量的嗜锇颗粒是病态的特征。高感家系受害植株中未受害针叶中,叶绿体呈圆形或破裂为不规则形,基质片层和基粒片层呈模糊状态或消失成碎屑,空泡化加大;由叶绿体包围的淀粉粒缩小,高感家系受害植株中未受害针叶中的光合作用已无法进行。
Plants against insects is not a passive suffering,but through many types of morphological structures and physiological processes to make a positive response. Because the resistant tree about insects have unique advantages,which in recent years,people take attention to the selection of target pest species resistant. In this paper,branches and pine needles on the measurement of external characteristics,and the paraffin section method and the ultra-thin sectioning were studied to explain the relationship between organizational structure,ultrastructure of needles of the correlation and the pest resistance relations,the following conclusions:
1. No victimization,the branch of high resistance family is more than in the mid-resistance and high susceptible family,the main branches and offshoots of the susceptible family are wider than the high resistance family and mid-resistance family,which indicated that the main branch and offshoots are more vulnerable,the diameter is bigger. High susceptible family was significantly longer than the length of needles in the mid-resistance and high resistance family,that needle length was negatively correlated with insect resistance.the bulk density of high resistance family and mid-resistance family was significantly bigger than the high susceptible family,which indicated the greater bulk density,the stronger the resistance. Damage by the pest,the pine needle retention rate in high resistance family and mid-resistance family is higher than high susceptible family. Affected,except for the susceptible family 469 in the main branch and offshoots’s diameter smaller than the healthy group, other families is bigger than the healthy group. By pests, the high resistance family 330 and mid-resistance family 155 does not reduce the needle retention,other family down the needle retention rate, indicating the stronger the resistance,the smaller the value of needle retention decreased.
2. The organizational structure of needles with different resistant shows that in the unaffected pest damage condition,cuticle thickness,the thickness of the lower epidermis,the epidermis thickness and the thickness of epidermis and cuticle thickness,mesophyll thickness,number and diameter of resin ducts,Victoria tube diameter was positively correlated with resistance to insects, while the epidermal thickness,stomatal density and cortical thickness,has nothing to do with insect resistance. In the damaged plantcompared damaged needle with undamaged needle,the resistance families showed cuticle,lower epidermis,the epidermis and horny and upper epidermis significantly thickened or changed,and susceptible families showed highly significant thinner. None insect needles in the damaged pinus compared to needles in the undamaged pinus,resistance families have thicker cuticle, but high susceptible family has thinner cuticle. None insect needles in the damaged pinus depend on thicking cuticle to increase their resistance..
From the microstructure point of view,high resistant family have deposits there in damaged position,Epidermis,mesophyll cells,resin ducts and other organizational structure within the cortex were normal in undamaged position; the mid-resistant family has more epidermal-rupture than resistant, and Mesophyll cell boundaries is not obvious,but within the cortex,vascular structure are normal; and high susceptible family on the epidermal cells and mesophyll cell shape,constriction reduced or broken, resin canal deformation,shrinkage and constricted. At the same time,containing material in inner cortical cells reduced. Epidermal structure,resin ducts,inner cortex,vascular bundles did not significantly changed in different resistant family,but mesophyll cells in high suscept family dramaticly reduced .
3. By images from TEM electron microscope,the high resistant family’s chloroplast lamellae in damaged needle is not clearer than undamaged needle in the same tree,Golgi increased,a large number of small bubbles formed,and osmiophilic particles appeared. Damaged by the pest,parts of the chloroplast lamellae disappeared,a large number of osmiophilic material appeared,part of the chloroplast envelope is broken in the mid-resistant family,which indicates that mid-resistant family is more seriously broken than the high resistant. High susceptible family chloroplast lamellae disappeared completely,more chloroplast envelope broken,so that light reaction and dark reaction can not be normally carried. At the same time,many starch grains appeared in chloroplast,released into the the cytoplasm;swelling of starch granules may be related to the collapse of the chloroplast membrane. A large number of electron-dense cytoplasm of the emergence of the ball,it may be secrete toxins to destroy the normal metabolism in cells .
With undamaged needles in high resistant family damaged plant,chloroplast stroma lamellae and grana lamellae boundaries marked,stacked neatly,mitochondria,cytoplasmic structure of normal,but the chloroplast outer membrane of a small amount of osmiophilic particles existed; part of the starch granules increased and transported out of chloroplasts. In undamaged needles in mid-resistant family damaged plant,chloroplast deformation,lamellar part of the fuzzy,chloroplast envelope surrounded by a large number of osmiophilic particles,which is pathological features. In high susceptible family,the chloroplasts were round or irregular rupture,stroma lamellae and grana lamellae were fuzzy state or disappear into the debris,increased vacuolization; surrounded by the chloroplast starch grain narrow, indicating that high high susceptible family’s were unable to carry out photosynthesis.
1.未受害情况下,高抗家系的分枝数多于中抗家系和高感家系的分枝数,高感家系的主枝和侧枝均比高抗家系和中抗家系粗,由此推测主枝和侧枝越粗越容易受松突圆蚧为害。高感家系的针叶长度显著大于中抗家系和高抗家系,表明针叶长度与抗虫性呈负相关。高抗家系和中抗家系的针叶容重显著大于高感家系,表明容重越大,抗性越强。受松突圆蚧危害后,高抗家系和中抗家系的松针保有率均比高感家系的松针保有率高。受害后,除高感家系469的主枝和侧枝比未受害组细以外,其它家系受害后均比未受害时增粗。受虫害后,高抗家系330和中抗家系155的松针保有率没有减少外,其它家系的松针保有率下降,说明抗性越强,松针保有率下降值越小。
2.针叶组织构造比较结果表明,在未受松突圆蚧为害情况下,角质膜厚度、下表皮厚度、表皮层厚度和表皮及角质厚度厚度、叶肉组织厚度、树脂道数目和直径、维管束直径与抗虫性呈正相关,而上表皮厚度、气孔密度、内皮层厚度与抗虫性无关。受害植株中受害针叶与未受害针叶相比,具有抗性的家系受害针叶表现出角质膜、下表皮、表皮层和角质及上表皮显著增厚或不变化,而感虫家系表现为极显著变薄。未受害植株的针叶和受害植株中未受害针叶相比,具有抗性的家系受害植株中未受害针叶的角质膜表现出增厚,而感虫性家系角质膜变薄,受害植株中未受害针叶主要通过角质膜的增厚来阻碍松突圆蚧的入侵。
从显微结构上看,高抗家系松突圆蚧吸食处有堆积物出现,未受害处表皮结构、叶肉细胞、树脂道和内皮层等组织结构均正常;中抗家系表皮破裂处比高抗家系增多,且叶肉细胞分界不明显,而内皮层、维管束等结构正常;而高感家系上表皮细胞和叶肉细胞变形、缢缩或破裂,树脂道变形,内皮层缢缩且细胞内内含物减少。不同抗性家系受害植株中未受害针叶与未受害植株针叶的组织结构相比,表皮结构、树脂道、内皮层、维管束都未有变化,高感家系叶肉细胞中细胞器急剧减少细胞内有沉积物出现。
3.通过透射电镜拍摄的电镜图片可以看出,受松突圆蚧为害后,高抗家系受害针叶的叶绿体片层没有正常的清晰,高尔基体增多,小泡大量形成,并且有嗜锇颗粒存在,表明高抗家系进入病态。受松突圆蚧危害后,中抗家系马尾松叶绿体片层部分消失,嗜锇物质大量存在,部分叶绿体被膜破裂,表明中抗家系的受害程度比高抗家系加重。高感家系受害针叶的叶绿体片层彻底消失,较多的叶绿体被膜破裂,从而使光反应和暗反应无法正常进行。同时,很多淀粉粒撑破叶绿体被膜,游离到细胞质中,淀粉粒的膨大可能与叶绿体膜的崩溃有关。细胞质中大量电子致密小球的出现,有可能是松突圆蚧在吸食细胞液的同时,还分泌一些毒素来破坏细胞的正常代谢机制。
高抗家系受害植株中未受害针叶中,叶绿体基质片层和基粒片层分界明显,堆叠整齐,线粒体、细胞质等结构正常,但叶绿体外膜有少量的嗜锇颗粒存在;部分淀粉粒增大并运出叶绿体。中抗家系受害植株中未受害针叶中,叶绿体变形,片层部分模糊,叶绿体被膜周围有大量的嗜锇颗粒是病态的特征。高感家系受害植株中未受害针叶中,叶绿体呈圆形或破裂为不规则形,基质片层和基粒片层呈模糊状态或消失成碎屑,空泡化加大;由叶绿体包围的淀粉粒缩小,高感家系受害植株中未受害针叶中的光合作用已无法进行。
Plants against insects is not a passive suffering,but through many types of morphological structures and physiological processes to make a positive response. Because the resistant tree about insects have unique advantages,which in recent years,people take attention to the selection of target pest species resistant. In this paper,branches and pine needles on the measurement of external characteristics,and the paraffin section method and the ultra-thin sectioning were studied to explain the relationship between organizational structure,ultrastructure of needles of the correlation and the pest resistance relations,the following conclusions:
1. No victimization,the branch of high resistance family is more than in the mid-resistance and high susceptible family,the main branches and offshoots of the susceptible family are wider than the high resistance family and mid-resistance family,which indicated that the main branch and offshoots are more vulnerable,the diameter is bigger. High susceptible family was significantly longer than the length of needles in the mid-resistance and high resistance family,that needle length was negatively correlated with insect resistance.the bulk density of high resistance family and mid-resistance family was significantly bigger than the high susceptible family,which indicated the greater bulk density,the stronger the resistance. Damage by the pest,the pine needle retention rate in high resistance family and mid-resistance family is higher than high susceptible family. Affected,except for the susceptible family 469 in the main branch and offshoots’s diameter smaller than the healthy group, other families is bigger than the healthy group. By pests, the high resistance family 330 and mid-resistance family 155 does not reduce the needle retention,other family down the needle retention rate, indicating the stronger the resistance,the smaller the value of needle retention decreased.
2. The organizational structure of needles with different resistant shows that in the unaffected pest damage condition,cuticle thickness,the thickness of the lower epidermis,the epidermis thickness and the thickness of epidermis and cuticle thickness,mesophyll thickness,number and diameter of resin ducts,Victoria tube diameter was positively correlated with resistance to insects, while the epidermal thickness,stomatal density and cortical thickness,has nothing to do with insect resistance. In the damaged plantcompared damaged needle with undamaged needle,the resistance families showed cuticle,lower epidermis,the epidermis and horny and upper epidermis significantly thickened or changed,and susceptible families showed highly significant thinner. None insect needles in the damaged pinus compared to needles in the undamaged pinus,resistance families have thicker cuticle, but high susceptible family has thinner cuticle. None insect needles in the damaged pinus depend on thicking cuticle to increase their resistance..
From the microstructure point of view,high resistant family have deposits there in damaged position,Epidermis,mesophyll cells,resin ducts and other organizational structure within the cortex were normal in undamaged position; the mid-resistant family has more epidermal-rupture than resistant, and Mesophyll cell boundaries is not obvious,but within the cortex,vascular structure are normal; and high susceptible family on the epidermal cells and mesophyll cell shape,constriction reduced or broken, resin canal deformation,shrinkage and constricted. At the same time,containing material in inner cortical cells reduced. Epidermal structure,resin ducts,inner cortex,vascular bundles did not significantly changed in different resistant family,but mesophyll cells in high suscept family dramaticly reduced .
3. By images from TEM electron microscope,the high resistant family’s chloroplast lamellae in damaged needle is not clearer than undamaged needle in the same tree,Golgi increased,a large number of small bubbles formed,and osmiophilic particles appeared. Damaged by the pest,parts of the chloroplast lamellae disappeared,a large number of osmiophilic material appeared,part of the chloroplast envelope is broken in the mid-resistant family,which indicates that mid-resistant family is more seriously broken than the high resistant. High susceptible family chloroplast lamellae disappeared completely,more chloroplast envelope broken,so that light reaction and dark reaction can not be normally carried. At the same time,many starch grains appeared in chloroplast,released into the the cytoplasm;swelling of starch granules may be related to the collapse of the chloroplast membrane. A large number of electron-dense cytoplasm of the emergence of the ball,it may be secrete toxins to destroy the normal metabolism in cells .
With undamaged needles in high resistant family damaged plant,chloroplast stroma lamellae and grana lamellae boundaries marked,stacked neatly,mitochondria,cytoplasmic structure of normal,but the chloroplast outer membrane of a small amount of osmiophilic particles existed; part of the starch granules increased and transported out of chloroplasts. In undamaged needles in mid-resistant family damaged plant,chloroplast deformation,lamellar part of the fuzzy,chloroplast envelope surrounded by a large number of osmiophilic particles,which is pathological features. In high susceptible family,the chloroplasts were round or irregular rupture,stroma lamellae and grana lamellae were fuzzy state or disappear into the debris,increased vacuolization; surrounded by the chloroplast starch grain narrow, indicating that high high susceptible family’s were unable to carry out photosynthesis.
引文
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