CN114868760B - 6-磷酸-海藻糖的应用及提升普通菜豆产量和抗病性的培育方法 - Google Patents
6-磷酸-海藻糖的应用及提升普通菜豆产量和抗病性的培育方法 Download PDFInfo
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Classifications
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- A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
- A01N57/00—Biocides, pest repellants or attractants, or plant growth regulators containing organic phosphorus compounds
- A01N57/10—Biocides, pest repellants or attractants, or plant growth regulators containing organic phosphorus compounds having phosphorus-to-oxygen bonds or phosphorus-to-sulfur bonds
- A01N57/16—Biocides, pest repellants or attractants, or plant growth regulators containing organic phosphorus compounds having phosphorus-to-oxygen bonds or phosphorus-to-sulfur bonds containing heterocyclic radicals
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01G—HORTICULTURE; CULTIVATION OF VEGETABLES, FLOWERS, RICE, FRUIT, VINES, HOPS OR SEAWEED; FORESTRY; WATERING
- A01G22/00—Cultivation of specific crops or plants not otherwise provided for
- A01G22/40—Fabaceae, e.g. beans or peas
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01G—HORTICULTURE; CULTIVATION OF VEGETABLES, FLOWERS, RICE, FRUIT, VINES, HOPS OR SEAWEED; FORESTRY; WATERING
- A01G7/00—Botany in general
- A01G7/06—Treatment of growing trees or plants, e.g. for preventing decay of wood, for tingeing flowers or wood, for prolonging the life of plants
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- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
Landscapes
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- Environmental Sciences (AREA)
- Ecology (AREA)
- Wood Science & Technology (AREA)
- Botany (AREA)
- Engineering & Computer Science (AREA)
- Forests & Forestry (AREA)
- Biodiversity & Conservation Biology (AREA)
- Zoology (AREA)
- Plant Pathology (AREA)
- Pest Control & Pesticides (AREA)
- General Health & Medical Sciences (AREA)
- Agronomy & Crop Science (AREA)
- Dentistry (AREA)
- Health & Medical Sciences (AREA)
- Agricultural Chemicals And Associated Chemicals (AREA)
Abstract
本发明公开了6‑磷酸‑海藻糖的应用及提升普通菜豆产量和抗病性的培育方法，通过喷施T6P溶液显著提升普通菜豆籽粒产量和抗病性，喷施T6P溶液后的普通菜豆叶片可溶性糖类含量显著提升，促进了植株生长、发育和碳水化合物的合成，同时叶片光合效率显著提高，促进营养物质向籽粒转运；此外，T6P诱导菜豆根中POD、SOD、PAL活性和H2O2含量显著提升，接菌28d后发病级别仅为4.2。因此，本发明将为普通菜豆籽粒产量和镰孢菌枯萎病抗性的提升提供一种新的方法，对改进我国普通菜豆产量和抗性水平有重要意义。
Description
技术领域
本发明涉及农作物培育技术领域，特别是一种6-磷酸-海藻糖的应用及提升普通菜豆产量和抗病性的培育方法。
背景技术
作物病害严重影响作物生长，造成产量损失，并威胁粮食安全。抗病性强的作物品种可以有效的控制病害，然而高抗性经常以“牺牲”产量为代价(Brown,2003；Nelson etal.,2018)。因此，研发协同调控抗病性和产量的作物化控调节剂对于保障粮食作物安全生产具有重要意义。普通菜豆(Phaseolus vulgaris L.)是世界上种植面积最大的食用豆类，产量约占全球食用豆类总产量的50％，特别是在非洲和美洲的部分地区更是作为一种重要的食物来源被广泛种植(张赤红等,2005；Pérez-Vega et al.,2010；Schmutz et al.,2014)。产量和抗病性是普通菜豆育种中最受育种家关注的两个要素，其中籽粒产量性状是影响普通菜豆生产潜力的重要因素，尖镰孢菌菜豆专化型(Fusarium oxysporumf.sp.phaseoli,Fop)引起的普通菜豆镰孢菌枯萎病(Fusarium wilt)是造成普通菜豆严重减产的重要原因，该病害作为最为严重的一种真菌性土传病害，分布于世界各地菜豆种植区域，每年给农户造成重大经济损失。
糖是能量和细胞碳骨架的供体，也是调控生长发育的重要信号分子。近年来，植物中6-磷酸-海藻糖(T6P)被发现具有类似动物胰岛素的功能，T6P水平与糖水平高度正相关，被称作糖水平的指示表；同时，T6P还可通过促进源-库转运等形式反馈调节糖水平(Paulet al.,2017)。作为维持糖稳态的核心激素，T6P广泛参与了调控植物的生长发育与逆境响应等生理过程(O'Hara et al.,2013)。尤为重要的是，T6P具有极大的改良作物产量的潜力。在玉米中异源表达水稻6-磷酸-海藻糖磷酸酶基因OsTPP1可直接提升9-49％的产量(Nuccio et al.,2015)。直接喷施可吸收的T6P前体亦可使小麦增产20％(Griffiths etal.,2016)。张健研究员团队与胡培松院士团队合作首次揭示了T6P调控水稻碳源分配与籽粒产量的机制，同时也为作物高产遗传改良提供了新思路(Li et al.,2022)。
植物病原体侵染寄主的过程中诱导细胞内积累大量的Tre。芸苔疟原虫(Plasmodiophora brassicae)侵染拟南芥引起Tre在根和下胚轴中迅速积累(Brodmann etal.,2002)。P.brassicae的感染导致拟南芥体内海藻糖酶基因表达显著增强，并且根和下胚轴中海藻糖酶活性也明显升高。此外，施用外源Tre可以诱导小麦对白粉病原菌(Blumeria graminis)的抗性(Reignault et al.,2001；Renard-Merlier et al.,2007)。外源Tre的喷施导致植物T6P含量的积累(Schluepmann et al.,2004)，鉴于T6P在糖信号传导途径中的重要作用，寄主体内积累的Tre很大程度是由T6P水解产生。
糖分子是调控植物生长发育、产量、籽粒性状和抗性的重要信号分子。海藻糖及其衍生物也是广泛参与植物生长发育与逆境响应等生理过程。T6P既是调控植物籽粒产量的糖代谢信号分子，同时也是合成Tre的重要前体，而Tre也已证明是调控植物产量和抗病性的重要信号分子。
发明内容
本发明的目的是要探索海藻糖代谢途径对普通菜豆籽粒产量特性、可溶性糖含量、光合生理指及镰孢菌枯萎病抗性的影响，提供一种6-磷酸-海藻糖的应用及提升普通菜豆产量和抗病性的培育方法。
为达到上述目的，本发明是按照以下技术方案实施的：
本发明的第一个目的是提供了一种6-磷酸-海藻糖在提升普通菜豆产量和抗病性中的应用。
本发明的第二个目的是提供了一种提升普通菜豆产量和抗病性的培育方法，在普通菜豆植株开花后每隔5d对普通菜豆植株喷洒一次6-磷酸-海藻糖溶液，至普通菜豆植株进入成熟期为止；和/或对7日龄的普通菜豆植株幼苗每日喷洒一次6-磷酸-海藻糖溶液，连续处理7d。
进一步地，所述6-磷酸-海藻糖溶液的浓度为1mM。
与现有技术相比，本发明通过喷施T6P溶液显著提升普通菜豆籽粒产量和抗病性，喷施T6P溶液后的普通菜豆叶片可溶性糖类含量显著提升，促进了植株生长、发育和碳水化合物的合成，同时叶片光合效率显著提高，促进营养物质向籽粒转运；此外，T6P诱导菜豆根中POD、SOD、PAL活性和H2O2含量显著提升，接菌28d后发病级别仅为4.2。因此，本发明将为普通菜豆籽粒产量和镰孢菌枯萎病抗性的提升提供一种新的方法，对改进我国普通菜豆产量和抗性水平有重要意义。
附图说明
图1为6-磷酸-海藻糖对普通菜豆防御因子的影响。
图2为6-磷酸-海藻糖对普通菜豆枯萎病抗性的影响。
具体实施方式
为使本发明的目的、技术方案及优点更加清楚明白，以下结合实施例，对本发明进行进一步的详细说明。此处所描述的具体实施例仅用于解释本发明，并不用于限定发明。
以下实施例采用普通菜豆品种龙饭豆1号和英国红芸豆作为试验材料，每个小区5m2作为一个重复，每个处理各3个重复，用于调查菜豆籽粒产量相关性状；将感病基因型品种英国红芸豆种子播种于灭菌的营养土中，23～28℃条件下，每日补充12h光照，用于枯萎病抗性研究。供试菌株为尖镰孢菌菜豆专化型FOP-DM01菌株(课题组保存)。病原菌首先接种于PDA平板上，25～28℃培养7d，然后用于接种。
实施例1
普通菜豆幼苗采用喷雾法处理，以蒸馏水作为溶剂，配制1mM 6-磷酸-海藻糖溶液(T6P)，在菜豆开花后每隔5d喷洒整株植物1次，至植株进入成熟期为止。
实施例2
准备7日龄的菜豆幼苗，每日喷1次1mM T6P，连续处理7d，用于接种病原菌，接种方法参照薛仁风等(2018)。
简述如下:将玉米粉与蛭石按1:2(V/V)的比例制成混合物，取400mL混合物于1000mL三角瓶中，121℃,0.2MP条件下30min,灭菌2次。向玉米粉混合物中加入50mL灭菌蒸馏水，混合均匀后接入一皿病原菌，25～28℃条件下培养7～10d，每天摇晃三角瓶使病原菌生长均匀。将接种体与灭菌的营养土按照1:10的比例混合，取0.01g接种混合土悬浮于1mL灭菌水，用血球计数板计算接种混合土中病原菌的含量，使接种终浓度达到5.0×106cfu g-1，取50μL悬浮液按1:10、1:100比例稀释后涂布于酸性PDA培养基(pH:5.0)上，通过培养基上生长的菌落数目和相应的稀释倍数计算每克接种土中含有病原菌的量，进一步验证血球计数板计算接种浓度的准确性。将用0.5％NaClO溶液表面灭菌的菜豆种子播种于上述接种混合土中，23～28℃温室条件下，每日补充光照12h，光照强度约300μmol m-2s-1，4～5周后调查发病情况。
发病级别调查参考王述民等(2006)的普通菜豆枯萎病1～9级的病情分级标准，具体如下。
对比例1
与实施例1的不同在于，在菜豆开花后每隔5d对整株植物喷洒1次蒸馏水，至植株进入成熟期为止。
分别取实施例1和对比例1的普通菜豆植株成熟期的植株叶片进行糖含量测定和光合生理指标测定，具体测定方法如下：
糖含量测定
在普通菜豆成熟期采集植株叶片，每个处理取3次生物学重复；将采集样品置于-20℃条件下保存，用于糖类物质含量的检测。海藻糖含量测定参考Li等(2022)的方法。简述如下：100mg干燥叶片均质于5mL 80％(v/v)热乙醇20min并离心。上清液在80℃下干燥并重新溶解于蒸馏水。悬浮液依次在0.1M H2SO4和0.6M氢氧化钠中煮沸。最后，混合物用蒽酮和98％硫酸100℃条件下处理10min。溶液冷却后测量630nm的吸光值。
葡萄糖、果糖、蔗糖和淀粉含量的测定参考Xue等(2021)的方法。简述如下：约0.5g叶片样品经过10mL乙醇/水混合物80℃条件下萃取30min(80:20,v/v)。溶液过滤后用氮气干燥，滤液重新溶解于等体积0.1mM CaEDTA溶液，过滤后通过高效液相色谱仪分析葡萄糖、果糖、蔗糖含量；叶片样品于80％乙醇中研磨并在40℃下孵育18h，9000g离心10min。蒸发上清液至干燥，重新溶解于2mLH2O和0.5mL氯仿。3000g离心10min，将含淀粉沉淀在55℃水浴条件下干燥，在3mL H2O中复溶并煮沸2h使淀粉糊化。添加3mL 0.1M醋酸盐缓冲液(pH 4.5)和0.5mL淀粉葡萄糖苷酶，55℃下温浴24h。9000g离心10min，保留上清液采用葡萄糖氧化酶试剂盒用于葡萄糖含量测定。淀粉转化的葡萄糖值乘以0.9转化为淀粉含量值。
测定结果如表1所示，结果表明：在T6P诱导条件下，龙饭豆1号和英国红芸豆叶片中海藻糖、葡萄糖、果糖、蔗糖和淀粉的含量均显著高于对照，其中英国红芸豆海藻糖、葡萄糖、蔗糖和淀粉水平最高，分别达37.4nmol/g、4.3μmol/g、5.5μmol/g和3.6μmol/g。龙饭豆1号果糖含量最高，达到8.4nmol/g。研究结果表明，T6P激活了普通菜豆叶片可溶性糖分子的合成，提升了植株糖类化合物的储备。
表1 6-磷酸-海藻糖对普通菜豆可溶性糖含量的影响
注：不同字母表示数据在P≤0.05水平上差异显著。
光合生理指标测定
在普通菜豆成熟期，检测植株叶片叶绿素总含量、净光合速率、气孔导度、胞间CO2浓度和蒸腾速率，每个处理取3次生物学重复，测定不同重复小区不同植株近同一部位的10张叶片。叶绿素总含量检测方法参考薛仁风等(2015)；净光合速率、气孔导度、胞间CO2浓度和蒸腾速率采用GB-1102便携式光合蒸腾仪进行测定。
测定结果如表2所示，结果表明：在T6P诱导条件下，龙饭豆1号和英国红芸豆叶片中叶绿素总含量略有升高，但变化并不明显。净光合速率、气孔导度、胞间CO2浓度和蒸腾速率均显著提升，其中龙饭豆1号净光合速率、胞间CO2浓度和蒸腾速率最高，分别达到18.6、249.6、6.5μmol/m2·s。英国红芸豆气孔导度最大，达0.57μmol/m2·s。研究结果表明，T6P促进普通菜豆叶片光合作用，为碳水化合物合成与积累奠定基础。
表2 6-磷酸-海藻糖对普通菜豆光合生理指标的影响
注：不同字母表示数据在P≤0.05水平上差异显著。
另外，实施例1中的采用浓度为1mM 6-磷酸-海藻糖(T6P)溶液从植株开花后定期喷施普通菜豆植株，成熟期调查植株籽粒产量相关性状的结果如表3所示，结果表明：在T6P诱导条件下，龙饭豆1号的单株荚数略有升高，但相比对照不显著，而单荚粒数、百粒重和理论产量均显著增加，分别达到4.5、42.7g和2250.5kg/hm2；而英国红芸豆但单株荚数、单荚粒数、百粒重和理论产量均明显著提高，分别达到25.1、5.3、43.2g和2549.6kg/hm2。研究结果表明，T6P激发了普通菜豆籽粒产量潜力，提升了籽粒产量相关性状。
表3 6-磷酸-海藻糖对普通菜豆籽粒性状的影响
注：不同字母表示数据在P≤0.05水平上差异显著
另外，取实施例2的普通菜豆植株分别进行植物防御关键因子、6-磷酸-海藻糖对普通菜豆抗病性的测定，具体测定方式如下：
分别在接种0、24和48h后的菜豆植株根部取样，每个处理取3次生物学重复；将采集样品置于-80℃条件下保存，用于过氧化物酶(POD)活性、超氧化物歧化酶(SOD)活性、丙氨酸解氨酶(PAL)活性和过氧化氢(H2O2)等防御反应因子的检测，方法参考Xue等(2021)。
POD活性测定
取菜豆根组织各100～150mg，分别加入预冷10mM磷酸缓冲液(137mM NaCl，2.7mMKC1，9.8mM Na2HPO4，1.7mM KH2PO4，pH:7.4)，充分匀浆，使各样品浓度均为0.1mg/μL，40℃条件下静置30min,13,000g离心20min。反应体系包括2.9mL含有1.25％(v/v)愈创木酚和0.1M H2O2的磷酸缓冲液，加入100μL酶粗提液，混合均匀后于25℃条件下反应5min，470nm波长下读取吸光值，测定POD活性。
SOD活性测定
取0.5g样品放入研钵中，加2mL 0.05M磷酸缓冲液(pH＝7.8)及少量石英砂，冰浴研磨，匀浆倒入10mL离心管中，再加3mL磷酸缓冲液冲洗研钵，4℃冷冻离心20min，上清液倒入试管中，置于0～4℃下保存待用。分别取Met溶液162mL，EDTA-Na2溶液0.6ml，磷酸缓冲液5.4mL，NBT溶液6mL，核黄素溶液6mL,混合后摇匀。分别取3mL反应混合液和30μL酶液于试管中，将试管置于光照培养箱中在4000lux光照下反应20min，同时做两支对照管，其中1支试管取3ml反应混合液加入30μL PBS(不加酶液)照光后测定作为最大光还原管，另1支只加缓冲液置于暗中测定时用于调零。以不照光的对照管(只有缓冲液并置于暗处)调零后，避光测560nm吸光值用于分析SOD活性。
PAL活性测定
将样品放入提前预冷的研钵中，加入液氮研磨至粉末。用50mM硼酸钠缓冲液(pH8.8，5mM巯基乙醇和1％PVP)溶解匀浆，使终浓度达到0.1mg/μL，将混合物在4℃下，12000rpm离心30min，取上清液测定酶活性。在0.1mL粗酶液中加入4mL硼酸缓冲液(50mM，pH8.8)，再加入1mL L-苯丙氨酸(20mM)，在37℃水浴60min。反应用0.2mL的3M HCl中止。以不加酶液为对照，在290nm下读取吸光值，测定PAL活性。
H2O2含量测定
100～150mg根组织样品加入预冷的0.01M磷酸缓冲液(pH:7.4)及5％三氯乙酸(TCA)的混和溶液(v/v，1/0.7)匀浆，使终浓度为0.1mg/μL，4℃条件下，10,000g离心10min。离心后取上清液，1.6mL上清液中加入0.4ml 50％TCA,0.4mL 10mM硫酸亚铁铵和0.2mL2.5M硫氰酸钾，混合溶液离心后在测定480nm吸光值，分析H2O2含量。
6-磷酸-海藻糖对植物防御反应因子的影响如图1所示(图1中：A：POD活性；B：SOD活性；C：PAL活性；D：H2O2含量，不同字母表示数据在P≤0.05水平上差异显著)，图1结果表明：接种病原菌24h后，T6P诱导的英国红芸豆根中重要防御因子POD活性、SOD活性、PAL活性和H2O2含量均显著提升，接菌48h后，菜豆根中防御因子活性或含量达到最高，POD活性、SOD活性和PAL活性分别达到78.3、32.9、0.054nk at/mg，H2O2含量达到0.33ng/mg。植物防御因子的激活显著提升了寄主对枯萎病原菌的抗性，为利用T6P进一步开展普通菜豆枯萎病抗性改良相关研究提供了理论依据。
6-磷酸-海藻糖对普通菜豆抗病性的影响如图2所示(不同字母表示数据在P≤0.05水平上差异显著)，由图2可知，接种病原菌14d后，1mM T6P处理的普通菜豆英国红芸豆(感病材料)植株对枯萎病原菌抗性明显增强；接种病原菌28d后，对照植株完全枯萎、死亡，而经过T6P溶液处理的植株发病级别显著低于对照植株，发病级别仅为4.2。
本发明将为普通菜豆籽粒产量和镰孢菌枯萎病抗性的提升提供一种新的方法，对改进我国普通菜豆产量和抗性水平有重要意义。
本发明的技术方案不限于上述具体实施例的限制，凡是根据本发明的技术方案做出的技术变形，均落入本发明的保护范围之内。
Claims (1)
1.6-磷酸-海藻糖在抗镰孢菌枯萎病性中的应用，其特征在于，在普通菜豆植株开花后每隔5 d对普通菜豆植株喷洒一次浓度为1mM的6-磷酸-海藻糖溶液，至普通菜豆植株进入成熟期为止；和/或对7日龄的普通菜豆植株幼苗每日喷洒一次浓度为1mM的6-磷酸-海藻糖溶液，连续处理7d。
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