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半夏对遮阴和高温响应机理的研究

Study on Responsing Mechanism to Shading and High Temperature Stress in Pinellia Ternate (Thunb.) Breit

【作者】 耿慧云

【导师】 王建华;

【作者基本信息】 山东农业大学 , 植物资源学, 2012, 博士

【摘要】 半夏(Pinellia ternate (thunb.) Breit.)为天南星科半夏属植物,以干燥块茎入药,属大宗中药材,需求量较大。为满足需求,各地开始引种栽培,但人工栽培长期以来产量低而不稳。半夏栽培研究越来越受重视。本实验以三个不同生态类型产区(江苏省泰州市、山东省潍坊市、河北省安国市)来源的栽培半夏为材料,在同一环境条件下栽培,研究半夏对遮阴和高温胁迫的生理响应,及遮阴对半夏块茎产量和总生物碱含量的影响。遮阴试验设置四个遮阴处理:0%、30%、50%、70%,遮阴度通过不同密度遮阳网调节,将三来源半夏种植于同一试验地。高温试验设置四个温度处理(昼/夜):25℃/20℃、30℃/20℃、35℃/20℃、40℃/20℃,材料种植于苗木育苗盘中,待叶片展开后移入光照培养箱进行温度处理。主要结论:1.随着遮阴程度的增加,半夏倒苗期持续时间增长。不同来源半夏倒苗持续时间亦有差异,山东和江苏半夏的持续时间较短,河北半夏的较长。2.遮阴显著降低了半夏的总产量,但有利于半夏珠芽的产生。随着遮阴程度的增加,一级块茎所占比例显著降低,四级块茎所占比例显著增加。三来源半夏的产量山东>江苏>河北,差异显著。半夏在同一地区连续种植存在品种退化现象,引种时应引用地理生态环境相近产区的半夏资源,以免造成减产。3.在0%-50%遮阴处理下,半夏叶片净光合速率日变化呈双峰曲线,有明显的“光合午休”;在70%处理下则呈单峰曲线,避免了“光合午休”。随着遮阴程度的增加,净光合速率日变化最大值降低且出现时间延迟。半夏叶片日光合值和日平均值随遮阴程度增加而降低。三来源半夏均有此表现。随着生育期的推进,光合速率随遮阴程度增加而降低。4.遮阴提高了叶片叶绿素及其组分的含量。随着光照强度的增加,半夏叶片出现光抑制,但这种抑制是可逆的,随着午后光照强度的降低,荧光参数均能恢复到早晨时的状态。5.遮阴降低了半夏叶片和块茎中可溶性糖和淀粉的含量,其关键酶活性也随遮阴程度增加而降低。同时遮阴处理下可溶性糖的合成和向块茎中的运送受到抑制,使得半夏块茎中淀粉含量降低,影响了半夏的产量形成。6.遮阴降低了半夏叶片和块茎中SOD、POD、CAT酶活性,第一次倒苗到第二次出苗间,处理间差异不显著,之前和之后处理间差异显著。生育前期,空气温度和土壤温度较高,随遮阴程度增加叶片和块茎中保护酶活性升高从而使MDA含量较低;生育后期,随遮阴程度增加MDA含量显著增加。7.遮阴显著提高了半夏块茎总生物碱含量,但总生物碱产量则随遮阴程度增加而显著减低。生物碱含量与块茎大小有负相关关系,其机理有待进一步研究。半夏总生物碱产量山东半夏>江苏半夏>河北半夏。8.随着胁迫温度的升高和时间的延长,半夏叶片的光合速率迅速降低。高温胁迫对光系统产生了不可逆的破坏,加速了叶片的衰老。高温显著降低了最大荧光、最大光化学效率、PSⅡ实际光化学效率。初始荧光、非光化学猝灭随着温度的升高先显著增加,后随叶片衰老而降低。三来源半夏均有此表现。9.高温胁迫破坏了半夏叶片和块茎中的保护酶系统。叶片中SOD、CAT酶活性随处理温度的升高和处理时间延长先升高后显著降低,POD酶活性则持续降低;块茎中保护酶活性则均随温度升高和处理时间延长先升高后降低,但块茎中酶活性降低的时间晚于叶片中的。MDA含量随温度升高和处理时间延长而显著增加,前期增加幅度小于后期。三来源半夏均有此表现。10.对照处理下半夏叶片和块茎中淀粉含量变化不显著,高温胁迫下淀粉含量显著降低。可溶性糖含量随着胁迫时间的延长先增加后显著降低。高温胁迫下SS、SPS、SSS酶失活,活性显著降低。高温胁迫降低了同化物的积累,从而加剧了其消耗。三来源半夏均有此表现。

【Abstract】 Pinellia ternate (thunb.) Breit. is herbaceous monocots of Aareeae Pnielll’a. It is one ofthe numeorus traditional Chinese l herbs,with its tubers used for medicine and there is ahigher requirement for it in recent years.Men has cultivated Pinellia ternate (thunb.) Breit.for30years.The growth of Pinellia ternate (thunb.) Breit. is greatly affected by cultivationconditions.In addition, the cost of cultivation is high and many diseases affect the tuber yieldof Pinellia ternate (thunb.) Breit. The research objects in the paper are Pinellias from threedifferent ecological production areas (Taizhou of Jiangsu, Weifang of Shandong, Anguo ofHebei), cultivated under the same environmental condition.Studies were taken that thePinellia’s physiological response to shading and high tempreture stress,and that the effect onmedicinal ingredients of Pinellia ternata Breit. tubers of shading conditions. Shading test setfour shading degrees, as follows:0%,30%,50%,70%, adjusting with different shade nets ofdifferent densities. The research materials were cultivated under the same environmental plots.High temperature test set four temperature treatments (day/night):25°C/20°C,30°C/20°C,35°C/20°C,40°C/20°C.The materials were planted in a seedling nursery andmoved into light incubato until the leaf unfolding for tempreture treatments.The main resultsare as follows:1.With the increasing of shading degree, the duration of sprout tumble of the Pinelliaternate (thunb.) Breit. become longer. The result was different in three P. ternata.s fromdifferent areas.The duration of P. ternata.from Shandong and Jiangsu was shorter,and the theduration of P. ternata.from Hebei was longer.Shade significantly reduced the total output of Pinellia, but it is conducive to thegeneration of Pinellia bulbils. With the increase of shade, the proportion of first level tuberwas significantly lower, fourth level tubers significantly increased.The output order of Pinelliafrom three sources was Shandong,Jiangsu,Hebei;and the difference was significant. Plantingin the same area continuously cause the degradation of the species. In order to avoidcuts,Breit in producing areas of the similar geographical environment should be selected forthe introduction.In0%-50%relatively light intensity conditions, the curve of leaf net photosynthetic dailyvariation of Pinellia ternata(Thunb.) Breit had two peaks and obvious phenomena of middaydepression.In70%relatively light intensity conditions, the diurnal net photosynthetic rate(NPR) exhibited a single peak curve to avoid phenomena of midday depression. Thegreater degree of shading,the maximum of diurnal net photosynthetic rate(NPR) was lowerwith time delay. Leaf total Pn/day and mean Pn/day of Pinellia ternata(Thunb.) Breit wassmaller with degree of shading increaseing. Pinellia ternate (Thunb.) Breit from three sourceshad the same result. With the growth of plant, the photosynthetic rate was lessening with thedegree of shading increasing.Shade could improve total chlorophyll content and the value of chlorophyll component.The greater degree of shading,photoinhibition had appeared in Pinellia ternate (Thunb.) Breitleaf. But this photoinhibition was reversible, the state of Chlorophyll fluorescence parametersrestored to its former good condition in the morning with reducing degree of shading aftermidday.Shading can reduce the content of soluble sugar and starch in leaves and tubers. And thekey enzyme’s activity also decreased along with the increase of shading degree. At the sametime, the synthesis of soluble sugar and the transport to the tubers would be restrained. So itreduced the starch content and influenced the yield.Shading reduced the SOD、POD and CAT enzymes’ activity of leaves and tubers. Therewas no significant difference in the treatments between the first sprout tumble and the secondtime. In the early time of the growth period, the air temperature and soil temperature werehigher. With the more shading degree leaves and tubers’ protection enzymes’ activity wouldincrease and the MDA content was lower. And in the later time of the growth period with themore shading degree the MDA content was higher.Shading increased the tuber’s total alkaloid content significantly, but the total alkaloidyield decreased along with the increase of shading degree. Alkaloid content and tuber’s sizehave negative correlation and its mechanism needs further research. Alkaloid yield in thethree places were the relationship that Shandong> Jiangsu> Hebei.2.High temperature stress significantly reduced the photosynthetic rate of Pinellia;withthe increase of temperature stress and the extension of time, leaf photosynthetic rate rapidlyreduced. High temperature reduced the content of chlorophyll, exacerbated the decompositionof chlorophyll, caused the light system damage irreversible, leaf senescence accelerating. Thehigh temperature significantly reduced the initial fluorescence,maximal fluorescence,maximum photochemical efficiency and PS Ⅱactual photochemical efficiency.Non-photochemical quenching increased significantly with temperatures rising and thendeclined with leaf senescence. High temperature stress destroyed protective enzyme system of the leaves and the tubers.The SOD and CAT enzymes’ activity in the leaves would increase at first and thensignificantly reduce with the increasing of the processing temperature and time but the PODenzyme’s activity continued to decrease. The protection enzymes’ activity of the tubers wouldincrease at first and then reduce with the increasing of the processing temperature and time.But the activity of the enzyme was reducing later in tubers than in leaves. MDA’s contentincreased significantly with the processing temperature and time. The increasing rate was lessin the early time than late. The there sources were all having this performance.High temperature significantly reduced the starch content in the Pinellia leaves andtubers.soluble sugar content increased first and then reduced significantly and SS, SPS, SSSactivity significantly reduced. High temperature stress reduces the accumulation ofassimilates, exacerbating their consumption, resulting in lower yields.

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