自然感染松材线虫后黑松与马尾松的病理生理学研究
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摘要
松材线虫病(Pine wilt disease)又称松树萎焉病或松树枯萎病,是松树的一种毁灭性病害。其病原——松材线虫Bursaphelenchus xylophilus原产北美,但该病害首先在日本发现并暴发成灾,目前已知分布于美国、加拿大、墨西哥、葡萄牙、西班牙韩国及中国等国家。我国自上世纪80年代初发现该病以来,扩散蔓延迅速,已对我国的松林生态系统构成了严重的破坏和威胁,成为我国最危险的森林生物灾害之一。该病害的发生涉及寄主树种、传播媒介、松材线虫与其携带的微生物、以及环境因素等多个方面,至今其致病机理尚存争议。
本文在松材线虫病疫区根据感病松树针叶变色程度及流脂状况,结合线虫分离,分别选取不同感病阶段的黑松(Pinus thunbergii)和马尾松(P. massoniana)为对象,研究了松材线虫自然侵染黑松和马尾松后,两种松树的光合作用及其相关生理生化变化过程。主要结果如下：
1.测定了两种松树不同感病阶段的净光合速率(Pn),发现黑松、马尾松在饱和光强下净光合速率均在感病初期表现最高，即在感病初期,两者Pn值反而大于健康阶段数值；之后随病害发展开始逐渐下降,至感病后期表现为负值：表明松材线虫的侵染严重影响了黑松、马尾松的光合作用,同时在线虫侵染初期,寄主植物都表现出一定的防御反应。另由叶绿素荧光参数分析可看出,光合速率的下降是由于光合系统Ⅱ反应中心实际进行光化学反应的效率(φPⅡ)下降引起的,而φPⅡ的降是由于PⅡ反应中心捕获光能效率(Fv'/Fm')下降和开放的PSⅡ中心数日(qP)下降共同导致。
2.两种松树在感病进程中,针叶叶绿素含量和类胡萝卜素含量在感病初期较健康阶段略有上升,之后整体呈下降趋势；此外,针叶含水量也表现出与之类似的变化趋势,认为这是引起松树萎蔫、叶绿素含量下降和光合作用速率下降的主要原因之同时也说明植物对病害胁迫表现出一定的补偿机制。实验表明,随着松材线虫侵染的加重,植株体内与抗性相关的抗氧化酶系统活性也会发生变化。两种松树感病后过氧化氢酶(CAT)活性即开始下降,而过氧化物酶(POD)活性和超氧化物歧化酶(SOD)活性却呈现先上升后下降趋势：说明松材线虫的侵入破坏了黑松、马尾松的抗氧化酶系统,寄主植物虽在病害胁迫初期作出了相应的防御反应,但随着松材线虫的大量繁殖,寄主自身保护作用逐渐减弱或丧失。
3.测定了马尾松、黑松在自然感病状态下针叶内还原糖及矿质营养元素的变化趋势。结果表明,两种松树随病程发展针叶内还原糖及N元素含量均逐渐降低；其余矿质元素含量变化不一。表明由于线虫的侵入,破坏了植物的组织结构,影响了寄主的新陈代谢和物质循环,继而影响到对各种元素的吸收、转移及利用。
4.首次采用ASD野外光谱仪测量了黑松和马尾松在松材线虫自然侵染后不同感病阶段的反射光谱,分析了其中光谱特征参数及叶绿素变化情况。结果表明：针对松树不同的发病阶段,应选择不同的光谱范围进行分析。从而能更好地反映植株病变程度与其光谱曲线间的关系;中红外波段反射光谱曲线对松树发病初期有一定的指示作用：红边位置、绿峰高度、红谷反射率、红边斜率、水分胁迫波段反射率等参数的动态变化趋势与松材线虫侵染后两种松树的病害症状具对应性；随病害程度加深两种松树叶绿素含量逐渐降低,且与光谱特征参数之间存显著线性关系。
5.开展了马尾松、黑松在自然感病状态下枝条的组织病理学研究,结果发现,马尾松和黑松自感病初期即表现出病变特征,并随感病时间推移病变加剧。感病初期马尾松枝条皮层薄壁细胞的细胞壁开始木质化，伴有细胞破裂病融合形成空腔；至感病中期皮层薄壁细胞、周皮的栓内层细胞、以及韧皮部薄壁细胞等细胞均发生木质化,细胞内含物增多,形成层细胞也发生一定程度的木质化,树脂道因周围细胞破裂而变形：至感病末期,木质部以外所有部分完全被术质化、大量细胞破裂。黑松发病症状与马尾松大致相同,但黑松发病时间相对稍晚,发病程度也较轻,这可能是因为两者对松材线虫病的抗性强弱有差异。比较分析寄主同一发病阶段上、中、下三个冠层中的组织切片,同一发病阶段植株组织病理学特征在空间上受危害程度也略存有差异,认为与松材线虫入侵的位置及在树体内的分布和数量有关。
Pine wilt disease is a fatal one caused by infection of the pine wilt nematode Bursaphelenchus xylophilus. The disease was first identified in Japan, where an outbreak occurred, and has been since spread to many countries including USA, Canada. Mexico. Portugal, South Korea and China. The pathogenesis of pine wilt is not clear, because it is affected by multiple factors including the plant hosts, longhorned beetles, nematodes and the microbes associated with them, as well as abiotic environmental factors. Pine wilt has posed enormous pressure on the ecosystem and caused huge economic loss in the disease epidemic countries. Since the beginning of the1980s', pine wilt has emerged and became widespread in China. It has been continuously threatening and destroying the pine ecosystem, and is one of the most dangerous biological disasters in the forest in China.
In this study, from a pine wilt epidemic area, we selected naturally infected Japanese black pine s (Pinus thunbergii) and Masson pines (P. massoniana) at different disease stages, isolated the nematode pathogen. and studied the host changes in photosynthesis and other physiological characteristics including color changes of needles and resinosis. The results are listed as follows:
1. Under saturating light intensity, both of the two pine species had the highest net photosynthetic rate (Pn) at the onset of the disease (higher than those of healthy pines). and the pines showed certain extent of defense responses. With disease progression, the Pn of the two species decreased to negative values, indicating that nematode infection interfered with the photosynthesis of Japanese black pines and Masson pines. In addition, analysis of chlorophyll fluorescence indicated that the decreases in photosynthetic rates were due to the decrease in the efficiency of photochemical reactions (ΦPSII) in photosystem Ⅱ, which was caused by the declines in the efficiency of light harvesting (Fv'/Fm') and the number of active PSII centers (qP).
2. During the onset of disease, the chlorophyll and carotenoid contents, as well as the water content in the needles of the two pine species slightly increased, and then dropped, which caused wilt and decrease in photosynthetic rate. In the diseased pines, the activity of catalase (CAT) reduced, and the activities of peroxidase (POD) and superoxide dismutase (SOD) also deceased eventually after slight increases at the beginning of the disease development. These changes indicated that nematode infection destroyed the antioxidant enzyme systems of Japanese black pines and Masson pines. Although the host plants had compensation mechanisms in response to stress in the early stage of the disease, the self-protection function was diminished and lost ultimately with the substantial proliferation of the pine wilt nematodes.
3. We have also analyzed the contents of reducing sugar and mineral nutrients in the diseased pine conifers. With disease progression, the contents of reducing sugar and nitrogen element decreased in both Japanese black pine and Masson pine. The changes in other minerals were inconsistent between the two types of pines. These results indicated that the structural damage caused by nematode infection significantly compromised metabolism, and affected the absorption, movement and utilization of different nutrients in the host plants.
4. We have measured the reflective spectrum of the Japanese black pines and Masson pines infected by pine wilt nematodes using an ASD field spectrometer, and analyzed the spectral characteristic parameters and the changes in chlorophyll content. To better understand the relationship between the disease severity and reflective spectrum, different wavelengths were used for the measurement of pines that were infected for various time periods. Reflective spectrum curves obtained with wavelength in the mid-infrared range were useful in indicating the onset of pine wilt disease. Specifically, the changes in parameters such as position of red edge, the height of green peak, the red valley reflectivity, the slope of red edge, and water stress band reflectivity were consistent with the disease symptoms of the two types of pines. The chlorophyll contents in the two pine species decreased with disease progression, and were significantly and linearly correlated with the spectral characteristic parameters.
5. Histopathological analyses showed typical lesions on the branches of Japanese black pine and Masson pine throughout the disease progression. During the early stages of disease development. the cell walls of cortical parenchyma cells on branches of Masson pines started to lignify. which was accompanied by cell ruptures and fusions to form cavities. During the middle stages of disease development, the cell walls of cortical parenchyma cells, periderm phelloderm cells, and phloem parenchyma cells also lignified. There was an increase in cell contents, and lignification was also observed in cambium cells. The resin canal deformed due to the rupture of surrounding cells. At the late stages of disease development, the majority of tissues outside the xylem were completely lignified and a large number of cells had ruptured. Japanese black pine and Masson pine had similar disease symptoms. Compared to Masson pine, the disease onset was delayed in Japanese black pine, and the disease was less severe. This might be due to the difference in resistance of the two different pine species to pine wilt disease. There was also difference in histopathology in sections obtained from upper, middle and lower canopies of the host plants at the same disease stage, which might be related to the differences in infection location, as well as the number and distribution of pine wood nematodes in the plants.
本文在松材线虫病疫区根据感病松树针叶变色程度及流脂状况,结合线虫分离,分别选取不同感病阶段的黑松(Pinus thunbergii)和马尾松(P. massoniana)为对象,研究了松材线虫自然侵染黑松和马尾松后,两种松树的光合作用及其相关生理生化变化过程。主要结果如下：
1.测定了两种松树不同感病阶段的净光合速率(Pn),发现黑松、马尾松在饱和光强下净光合速率均在感病初期表现最高，即在感病初期,两者Pn值反而大于健康阶段数值；之后随病害发展开始逐渐下降,至感病后期表现为负值：表明松材线虫的侵染严重影响了黑松、马尾松的光合作用,同时在线虫侵染初期,寄主植物都表现出一定的防御反应。另由叶绿素荧光参数分析可看出,光合速率的下降是由于光合系统Ⅱ反应中心实际进行光化学反应的效率(φPⅡ)下降引起的,而φPⅡ的降是由于PⅡ反应中心捕获光能效率(Fv'/Fm')下降和开放的PSⅡ中心数日(qP)下降共同导致。
2.两种松树在感病进程中,针叶叶绿素含量和类胡萝卜素含量在感病初期较健康阶段略有上升,之后整体呈下降趋势；此外,针叶含水量也表现出与之类似的变化趋势,认为这是引起松树萎蔫、叶绿素含量下降和光合作用速率下降的主要原因之同时也说明植物对病害胁迫表现出一定的补偿机制。实验表明,随着松材线虫侵染的加重,植株体内与抗性相关的抗氧化酶系统活性也会发生变化。两种松树感病后过氧化氢酶(CAT)活性即开始下降,而过氧化物酶(POD)活性和超氧化物歧化酶(SOD)活性却呈现先上升后下降趋势：说明松材线虫的侵入破坏了黑松、马尾松的抗氧化酶系统,寄主植物虽在病害胁迫初期作出了相应的防御反应,但随着松材线虫的大量繁殖,寄主自身保护作用逐渐减弱或丧失。
3.测定了马尾松、黑松在自然感病状态下针叶内还原糖及矿质营养元素的变化趋势。结果表明,两种松树随病程发展针叶内还原糖及N元素含量均逐渐降低；其余矿质元素含量变化不一。表明由于线虫的侵入,破坏了植物的组织结构,影响了寄主的新陈代谢和物质循环,继而影响到对各种元素的吸收、转移及利用。
4.首次采用ASD野外光谱仪测量了黑松和马尾松在松材线虫自然侵染后不同感病阶段的反射光谱,分析了其中光谱特征参数及叶绿素变化情况。结果表明：针对松树不同的发病阶段,应选择不同的光谱范围进行分析。从而能更好地反映植株病变程度与其光谱曲线间的关系;中红外波段反射光谱曲线对松树发病初期有一定的指示作用：红边位置、绿峰高度、红谷反射率、红边斜率、水分胁迫波段反射率等参数的动态变化趋势与松材线虫侵染后两种松树的病害症状具对应性；随病害程度加深两种松树叶绿素含量逐渐降低,且与光谱特征参数之间存显著线性关系。
5.开展了马尾松、黑松在自然感病状态下枝条的组织病理学研究,结果发现,马尾松和黑松自感病初期即表现出病变特征,并随感病时间推移病变加剧。感病初期马尾松枝条皮层薄壁细胞的细胞壁开始木质化，伴有细胞破裂病融合形成空腔；至感病中期皮层薄壁细胞、周皮的栓内层细胞、以及韧皮部薄壁细胞等细胞均发生木质化,细胞内含物增多,形成层细胞也发生一定程度的木质化,树脂道因周围细胞破裂而变形：至感病末期,木质部以外所有部分完全被术质化、大量细胞破裂。黑松发病症状与马尾松大致相同,但黑松发病时间相对稍晚,发病程度也较轻,这可能是因为两者对松材线虫病的抗性强弱有差异。比较分析寄主同一发病阶段上、中、下三个冠层中的组织切片,同一发病阶段植株组织病理学特征在空间上受危害程度也略存有差异,认为与松材线虫入侵的位置及在树体内的分布和数量有关。
Pine wilt disease is a fatal one caused by infection of the pine wilt nematode Bursaphelenchus xylophilus. The disease was first identified in Japan, where an outbreak occurred, and has been since spread to many countries including USA, Canada. Mexico. Portugal, South Korea and China. The pathogenesis of pine wilt is not clear, because it is affected by multiple factors including the plant hosts, longhorned beetles, nematodes and the microbes associated with them, as well as abiotic environmental factors. Pine wilt has posed enormous pressure on the ecosystem and caused huge economic loss in the disease epidemic countries. Since the beginning of the1980s', pine wilt has emerged and became widespread in China. It has been continuously threatening and destroying the pine ecosystem, and is one of the most dangerous biological disasters in the forest in China.
In this study, from a pine wilt epidemic area, we selected naturally infected Japanese black pine s (Pinus thunbergii) and Masson pines (P. massoniana) at different disease stages, isolated the nematode pathogen. and studied the host changes in photosynthesis and other physiological characteristics including color changes of needles and resinosis. The results are listed as follows:
1. Under saturating light intensity, both of the two pine species had the highest net photosynthetic rate (Pn) at the onset of the disease (higher than those of healthy pines). and the pines showed certain extent of defense responses. With disease progression, the Pn of the two species decreased to negative values, indicating that nematode infection interfered with the photosynthesis of Japanese black pines and Masson pines. In addition, analysis of chlorophyll fluorescence indicated that the decreases in photosynthetic rates were due to the decrease in the efficiency of photochemical reactions (ΦPSII) in photosystem Ⅱ, which was caused by the declines in the efficiency of light harvesting (Fv'/Fm') and the number of active PSII centers (qP).
2. During the onset of disease, the chlorophyll and carotenoid contents, as well as the water content in the needles of the two pine species slightly increased, and then dropped, which caused wilt and decrease in photosynthetic rate. In the diseased pines, the activity of catalase (CAT) reduced, and the activities of peroxidase (POD) and superoxide dismutase (SOD) also deceased eventually after slight increases at the beginning of the disease development. These changes indicated that nematode infection destroyed the antioxidant enzyme systems of Japanese black pines and Masson pines. Although the host plants had compensation mechanisms in response to stress in the early stage of the disease, the self-protection function was diminished and lost ultimately with the substantial proliferation of the pine wilt nematodes.
3. We have also analyzed the contents of reducing sugar and mineral nutrients in the diseased pine conifers. With disease progression, the contents of reducing sugar and nitrogen element decreased in both Japanese black pine and Masson pine. The changes in other minerals were inconsistent between the two types of pines. These results indicated that the structural damage caused by nematode infection significantly compromised metabolism, and affected the absorption, movement and utilization of different nutrients in the host plants.
4. We have measured the reflective spectrum of the Japanese black pines and Masson pines infected by pine wilt nematodes using an ASD field spectrometer, and analyzed the spectral characteristic parameters and the changes in chlorophyll content. To better understand the relationship between the disease severity and reflective spectrum, different wavelengths were used for the measurement of pines that were infected for various time periods. Reflective spectrum curves obtained with wavelength in the mid-infrared range were useful in indicating the onset of pine wilt disease. Specifically, the changes in parameters such as position of red edge, the height of green peak, the red valley reflectivity, the slope of red edge, and water stress band reflectivity were consistent with the disease symptoms of the two types of pines. The chlorophyll contents in the two pine species decreased with disease progression, and were significantly and linearly correlated with the spectral characteristic parameters.
5. Histopathological analyses showed typical lesions on the branches of Japanese black pine and Masson pine throughout the disease progression. During the early stages of disease development. the cell walls of cortical parenchyma cells on branches of Masson pines started to lignify. which was accompanied by cell ruptures and fusions to form cavities. During the middle stages of disease development, the cell walls of cortical parenchyma cells, periderm phelloderm cells, and phloem parenchyma cells also lignified. There was an increase in cell contents, and lignification was also observed in cambium cells. The resin canal deformed due to the rupture of surrounding cells. At the late stages of disease development, the majority of tissues outside the xylem were completely lignified and a large number of cells had ruptured. Japanese black pine and Masson pine had similar disease symptoms. Compared to Masson pine, the disease onset was delayed in Japanese black pine, and the disease was less severe. This might be due to the difference in resistance of the two different pine species to pine wilt disease. There was also difference in histopathology in sections obtained from upper, middle and lower canopies of the host plants at the same disease stage, which might be related to the differences in infection location, as well as the number and distribution of pine wood nematodes in the plants.
引文
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