本文關(guān)鍵詞：金銀花對(duì)土壤干旱—復(fù)水的光合生理響應(yīng)　出處：《山東農(nóng)業(yè)大學(xué)》2017年博士論文　論文類型：學(xué)位論文

  更多相關(guān)文章： 金銀花 干旱脅迫 復(fù)水 光合作用 水分閾值

【摘要】：金銀花在沂蒙山區(qū)被廣泛用作水土保持造林和藥用經(jīng)濟(jì)栽培植物,為沂蒙山區(qū)的生態(tài)恢復(fù)和經(jīng)濟(jì)發(fā)展做出了卓越貢獻(xiàn)。但是沂蒙山區(qū)春夏季土壤重度干旱發(fā)生概率呈增大趨勢(shì),加之沂蒙山區(qū)干旱瘠薄的立地條件,增加了金銀花受到重度干旱脅迫的風(fēng)險(xiǎn),嚴(yán)重制約了金銀花的生長(zhǎng)發(fā)育和生產(chǎn)力。當(dāng)土壤干旱發(fā)生后,金銀花通過(guò)一系列生理生態(tài)學(xué)機(jī)制去抵御和適應(yīng)脅迫,金銀花光合生理對(duì)干旱脅迫的響應(yīng)過(guò)程與復(fù)水后的恢復(fù)過(guò)程相關(guān)聯(lián),因此需要通過(guò)量化其光合生理參數(shù)與土壤水分含量的關(guān)系來(lái)對(duì)金銀花外在形態(tài)特征和內(nèi)部生理機(jī)制的變化過(guò)程進(jìn)行深入的研究,以便對(duì)金銀花進(jìn)行科學(xué)高效的水分管理。但是目前關(guān)于金銀花光合生理對(duì)干旱脅迫響應(yīng)的研究尚少,關(guān)于重度土壤干旱脅迫對(duì)金銀花光合生理的影響以及復(fù)水后金銀花光合生理的恢復(fù)性尚未見報(bào)道,這些研究的缺乏不利于金銀花應(yīng)對(duì)干旱脅迫的水分管理措施體系的建立。本文以沂蒙山區(qū)常見的水土保持植物和經(jīng)濟(jì)樹種——金銀花一年生苗木為試驗(yàn)材料,于2016年3—10月在山東農(nóng)業(yè)大學(xué)林學(xué)試驗(yàn)站溫室內(nèi)進(jìn)行盆栽試驗(yàn),并結(jié)合實(shí)驗(yàn)室室內(nèi)分析試驗(yàn),進(jìn)行土壤水分的控制條件下金銀花光合作用氣體交換參數(shù)、葉綠素?zé)晒鈪?shù)、光譜特征參數(shù)和酶活性指標(biāo)的監(jiān)測(cè),以凈光合速率(Pn)、水分利用效率(WUE)為衡量土壤水分(RSWC)影響金銀花光合作用產(chǎn)量和效率的指標(biāo)(簡(jiǎn)稱為“產(chǎn)”、“效”),以光合作用氣孔調(diào)節(jié)機(jī)理、葉綠素?zé)晒庾兓瘷C(jī)理和葉片形態(tài)變化特征為依據(jù),分析了金銀花光合生理在“干旱脅迫—脅迫持續(xù)—復(fù)水”過(guò)程中的變化特征,探討了金銀花對(duì)連續(xù)多級(jí)土壤水分梯度、重度土壤干旱脅迫的光合生理響應(yīng)以及重度土壤干旱后復(fù)水金銀花光合生理的恢復(fù)性機(jī)理,并在此基礎(chǔ)上,得出了從土壤飽和含水量至Pn=0土壤含水量(凋萎含水量,此時(shí)金銀花開始萎蔫并逐漸凋落)范圍內(nèi)水分有效性分級(jí)體系和重度土壤干旱水分閾值(RSWCSL-NSL、RSWCPn=0),并根據(jù)金銀花在重度干旱下光合作用“產(chǎn)”、“效”降到最低值的速度(簡(jiǎn)稱為“速”)以及復(fù)水后金銀花光合作用“產(chǎn)”、“效”恢復(fù)到最大值的速度(“速”),得出了基于“速—產(chǎn)—效”金銀花響應(yīng)干旱脅迫光合作用降低及復(fù)水后光合作用恢復(fù)性的關(guān)鍵閾值,形成了金銀花在重度土壤干旱下光合作用降低及復(fù)水后光合作用恢復(fù)性的動(dòng)態(tài)過(guò)程體系,為重度土壤干旱下金銀花的水分管理工作提供科學(xué)依據(jù),而且本文研究中確定的不同水分閾值具有普遍的生理生態(tài)學(xué)意義,對(duì)其它植物也具有適用性。論文主要的結(jié)論如下:1)金銀花葉片光合作用氣體交換參數(shù)、葉綠素?zé)晒鈪?shù)、光譜特征參數(shù)和酶活性指標(biāo)對(duì)土壤水分含量(RSWC)具有閾值響應(yīng)特征,土壤水分過(guò)高或過(guò)低都會(huì)影響金銀花光合作用�；诮疸y花光合作用受抑制的特征將金銀花從土壤飽和含水量至Pn=0含水量之間土壤水分范圍按照水分脅迫程度劃分為適宜金銀花生長(zhǎng)的水分范圍(70.54%≤RSWC≤83.50%)、輕度水分脅迫水分范圍(55.07%≤RSWC70.54%或RSWC83.50%)、中度水分脅迫水分范圍(29.71%RSWC55.07%)、重度水分脅迫水分范圍(RSWC≤29.71%)。以Pn和WUE為表征金銀花光合作用“產(chǎn)”和“效”的指標(biāo),應(yīng)用臨界值分類法將土壤水分梯度分為“無(wú)產(chǎn)無(wú)效水”(RSWC≤13.82%)、“低產(chǎn)低效水”(13.82RSWC≤29.71%)、“中產(chǎn)中效水”(29.71%RSWC≤30.64%或RSWC93.39%)、“中產(chǎn)高效水”(30.64%RSWC≤51.53%)、“高產(chǎn)中效水”(83.76RSWC≤93.39)和“高產(chǎn)高效水”(51.53RSWC≤83.76%)6個(gè)等級(jí)(利用類型)。金銀花重度干旱水分范圍(RSWC≤29.71%)包括土壤水分有效性分級(jí)中的“低產(chǎn)低效水”和“無(wú)產(chǎn)無(wú)效水”,在該水分范圍內(nèi),以金銀花氣孔機(jī)理轉(zhuǎn)折水分點(diǎn)(RSWCSL-NSL)和Pn為0水分點(diǎn)(RSWCPn=0,即金銀花葉片萎蔫水分點(diǎn))作為金銀花重度干旱脅迫的關(guān)鍵水分閾值。2)金銀花對(duì)土壤水分脅迫的響應(yīng),同時(shí)與土壤水分脅迫程度及其脅迫持續(xù)時(shí)間有關(guān)。在重度土壤干旱各水分閾值(即水分閾值1:RSWCSL-NSL,水分閾值2:RSWCPn=0)及其脅迫持續(xù)時(shí)間(30d)耦合作用下,金銀花凈光合速率(Pn)、水分利用效率(WUE)、蒸騰速率(Tr)、氣孔導(dǎo)度(Gs)、氣孔限制值(Ls)、表觀光合量子效率(Φ)、最大凈光合速率(Pnmax)、光飽和點(diǎn)(LSP)、暗呼吸速率(Rd)、最大熒光(Fm)、PSⅡ最大光化學(xué)效率(Fv/Fm)、PSⅡ?qū)嶋H光化學(xué)效率(ΦPSⅡ)、表觀光合電子傳遞速率(ETR)、葉綠素(Chl)熒光光化學(xué)淬滅(qP)、非光化學(xué)淬滅(NPQ)、水分指數(shù)(WI)、葉綠素歸一化指數(shù)(Chl NDI)、歸一化顏色指數(shù)(NDCI)以及超氧化物歧化酶(SOD)、過(guò)氧化物酶(POD)、過(guò)氧化氫酶(CAT)的酶活性顯著降低,胞間CO2濃度(Ci)、光補(bǔ)償點(diǎn)(LCP)、初始熒光(Fo)、光化學(xué)反射指數(shù)(PRI)、丙二醛(MDA)含量顯著上升,導(dǎo)致金銀花“產(chǎn)”、“效”下降的主要原因是非氣孔限制,干旱脅迫期間金銀花葉片出現(xiàn)萎蔫并逐漸凋落,葉片光合機(jī)構(gòu)受到不可逆破壞,金銀花光合產(chǎn)量嚴(yán)重降低。確定了水分閾值1(RSWCSL-NSL)、水分閾值2(RSWCPn=0)分別持續(xù)25d和5d的干旱脅迫為金銀花基于“速—產(chǎn)—效”光合作用降低的水分閾值(即閾值1(25)、閾值2(5)),干旱脅迫重于此閾值時(shí),Pn、WUE將達(dá)到最低值(0值)。3)金銀花復(fù)水后光合作用的恢復(fù)性(恢復(fù)程度和恢復(fù)速度)同時(shí)與復(fù)水前重度土壤干旱不同水分閾值及其脅迫持續(xù)時(shí)間有關(guān),而且金銀花在重度土壤干旱后光合作用的降低和復(fù)水后光合作用的恢復(fù)性緊密聯(lián)系。金銀花在不同水分閾值(RSWCSL-NSL、RSWCPn=0)處分別進(jìn)行持續(xù)脅迫后復(fù)水其氣體交換參數(shù)、葉綠素?zé)晒鈪?shù)、光譜特征參數(shù)和酶活性指標(biāo)在復(fù)水的前15d左右恢復(fù)相對(duì)較快,之后或者達(dá)到穩(wěn)定狀態(tài)或者繼續(xù)緩慢恢復(fù)。根據(jù)金銀花復(fù)水后光合生理的恢復(fù)性,將其在水分閾值2持續(xù)脅迫時(shí)的萎蔫情況分為臨時(shí)萎蔫,即水分閾值2(RSWCPn=0)持續(xù)0d(閾值2(0))時(shí)的萎蔫;以及永久萎蔫,即水分閾值2(RSWCPn=0)持續(xù)10d以上(即閾值2(10)、閾值2(20)、閾值2(30))。以金銀花“產(chǎn)”、“效”最高的水分處理(RSWC=79.59%)的光合生理參數(shù)平均值(PAR=1200μmol·m-2·s-1)為對(duì)照,以所有水分處理(土壤飽和含水量到Pn=0含水量)的光合生理參數(shù)平均值(PAR=1200μmol·m-2·s-1)為均值,對(duì)各復(fù)水處理光合生理參數(shù)的恢復(fù)效果進(jìn)行評(píng)價(jià)。根據(jù)金銀花在重度土壤干旱脅迫后復(fù)水光合作用基于“速—產(chǎn)—效”的恢復(fù)性,確定了RSWCSL-NSL持續(xù)0d的干旱脅迫(即閾值1(0))為金銀花光合生理恢復(fù)性的第1級(jí)水分閾值,干旱脅迫輕于此閾值,復(fù)水15天內(nèi)Pn、WUE恢復(fù)到甚至超過(guò)對(duì)照水平;以RSWCSL-NSL持續(xù)10d、RSWCPn=0持續(xù)0d的干旱脅迫(即閾值1(10)、閾值2(0))為恢復(fù)性的第2級(jí)水分閾值,干旱脅迫輕于此閾值而重于第1個(gè)閾值時(shí),復(fù)水25d內(nèi)Pn、WUE恢復(fù)至均值以上水平;以RSWCSL-NSL持續(xù)20d、RSWCPn=0持續(xù)10d的干旱脅迫(即閾值1(20)、閾值2(10))為恢復(fù)性的第3級(jí)水分閾值,當(dāng)干旱脅迫重于此閾值時(shí),復(fù)水到30d時(shí),Pn、WUE都沒有恢復(fù)到均值水平。
[Abstract]:Honeysuckle is widely used in Yimeng mountain afforestation and medicinal plant cultivation economy of soil and water conservation, made outstanding contributions to the ecological restoration of Yimeng mountain area and economic development. But the Yimeng mountain spring and summer soil increases the probability of occurrence of severe drought, drought and barren in Yimeng mountain area and site conditions, increasing the risk of severe drought stress by honeysuckle, serious restrict the growth and productivity of honeysuckle. When soil drought occurred, honeysuckle to resist and adapt to stress through a series of physiological and ecological mechanism, physiological recovery process of gold and silver money in response to drought stress and rewatering associated, so through the quantitative relationship between the photosynthetic physiological parameters and soil water content to change the honeysuckle external morphological characteristics and internal physiological mechanism of the process requires in-depth research, in order to in Honeysuckle The scientific water management efficiency. But the current research on the gold and silver money physiological response to drought stress is less about the reported recovery of severe effects of soil drought stress on Photosynthetic Physiology and all gold and silver after rehydration of gold and silver spend physioecological, the establishment of these studies lack is not conducive to the system of water management measures in response to the drought of honeysuckle stress. In this paper, the Yimeng mountain area common plant for soil and water conservation and economic species of honeysuckle seedlings as test materials, in 2016 3 to October at the Shandong Agricultural University Forestry Experimental Station in greenhouse pot experiment, combined with analysis of test laboratory, exchange parameters controlling conditions of soil moisture under honeysuckle photosynthesis chlorophyll fluorescence parameters of gas. Monitoring, spectral characteristic parameters and enzyme activity index, the net photosynthetic rate (Pn), water use efficiency (WUE) for The measure of soil moisture (RSWC) affect the yield and efficiency of photosynthesis index of honeysuckle (hereinafter referred to as the "production" and "efficiency"), stomatal regulation on the photosynthesis mechanism, chlorophyll fluorescence and leaf morphology change mechanism according to the variation characteristics, analysis of Photosynthetic Physiology in "all gold and silver drought - stress - complex variation of water" the study of honeysuckle on continuous multistage soil moisture gradient, severe soil drought stress on photosynthetic physiological response and severe soil drought rewatering after physiological recovery of gold and silver money mechanism, and on this basis, obtained from the soil saturated water content and Pn=0 of soil moisture (wilting, the honeysuckle began wilting and gradually fall within the scope of water availability) grading system and severe soil drought water threshold (RSWCSL-NSL, RSWCPn=0), and according to the degree of drought on Photosynthesis in heavy honeysuckle "Production" and "efficiency" to a minimum value of speed (referred to as the "speed") and after rewatering honeysuckle Photosynthesis "production", "effect" back to the maximum speed ("speed"), obtained based on the "speed - production effect of honeysuckle in response to drought stress reduced photosynthesis and water recovery after the recovery of photosynthesis of the critical threshold, the formation of honeysuckle in severe soil drought reduced photosynthesis and rehabilitation of water dynamic process of photosynthesis system recovery, and provide scientific basis for severe soil drought under honeysuckle water management work, and the research on different water threshold determined with universal significance of physiological ecology, it is applicable on the other plants. The main conclusions are as follows: 1) gas exchange parameters of honeysuckle leaf photosynthesis, chlorophyll fluorescence parameters, spectral characteristic parameters and enzyme activity index of soil moisture content (RSWC) with threshold response characteristics, soil moisture is too high or too low will affect the photosynthesis characteristics of honeysuckle honeysuckle. Photosynthesis inhibition based on honeysuckle from saturated soil water content and water content of Pn=0 soil moisture range according to the stress level is divided into the suitable moisture range of honeysuckle growth (RSWC = 70.54% ~ 83.50%), mild water water stress range (55.07% RSWC70.54% or RSWC83.50%), moderate water stress and water range (29.71%RSWC55.07%), severe water stress water range (RSWC < 29.71%). Pn and WUE for the characterization of honeysuckle Photosynthesis "production" and "efficiency" of the index, using the critical value method to soil moisture gradient is divided into "no property is invalid water" (RSWC 13.82%), "low efficiency water" (13.82RSWC = 29.71%), "the effect of middle water" (29.71%RSWC < 30.64% or RSWC93.39%), "middle high water" (30 .64%RSWC = 51.53%), "high efficiency water" (83.76RSWC = 93.39) and the "high water" (51.53RSWC 83.76%) 6 grades (by type). Honeysuckle severe drought water range (RSWC = 29.71%) including the classification of soil water availability in low yield water and no water production is invalid "in the moisture range, with honeysuckle stomatal mechanism turning moisture point (RSWCSL-NSL) and Pn 0 water points (RSWCPn=0, namely honeysuckle leaf wilting moisture point) as a key water threshold of severe drought stress.2 honeysuckle honeysuckle) response to soil moisture stress, and soil water stress degree and duration of stress in severe soil drought. The water threshold (i.e. water threshold 1:RSWCSL-NSL, water threshold 2:RSWCPn=0) and stress duration (30d) coupling, honeysuckle, net photosynthetic rate (Pn), water use efficiency (WUE), transpiration 閫熺巼(Tr),姘斿瓟瀵煎害(Gs),姘斿瓟闄愬埗鍊，

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