Tillering is an important agronomic trait of rice(Oryza sativa)that affects the number of effective panicles,thereby affecting yields.The phytohormone auxin plays a key role in tillering.Here we identified the high ti...Tillering is an important agronomic trait of rice(Oryza sativa)that affects the number of effective panicles,thereby affecting yields.The phytohormone auxin plays a key role in tillering.Here we identified the high tillering and semi-dwarf 1(htsd1)mutant with auxin-deficiency root characteristics,such as shortened lateral roots,reduced lateral root density,and enlarged root angles.htsd1 showed reduced sensitivity to auxin,but the external application of indole-3-acetic acid(IAA)inhibited its tillering.We identified the mutated gene in htsd1 as AUXIN1(OsAUX1,LOC_Os01g63770),which encodes an auxin influx transporter.The promoter sequence of OsAUX1 contains many SQUAMOSA PROMOTER BINDING PROTEIN-LIKE(SPL)binding sites,and we demonstrated that SPL7 binds to the OsAUX1 promoter.TEOSINTE BRANCHED1(OsTB1),a key gene that negatively regulates tillering,was significantly downregulated in htsd1.Tillering was enhanced in the OsTB1 knockout mutant,and the external application of IAA inhibited tiller elongation in this mutant.Overexpressing OsTB1 restored the multi-tiller phenotype of htsd1.These results suggest that SPL7 directly binds to the OsAUX1 promoter and regulates tillering in rice by altering OsTB1 expression to modulate auxin signaling.展开更多
Recently developed ‘super’ rice cultivars with greater yield potentials often suffer from the problem of poor grain filling, especially in inferior spikelets. Here, we studied the activities of enzymes related to st...Recently developed ‘super’ rice cultivars with greater yield potentials often suffer from the problem of poor grain filling, especially in inferior spikelets. Here, we studied the activities of enzymes related to starch metabolism in rice stems and grains, and the microstructures related to carbohydrate accumulation and transportation to investigate the effects of different water regimes on grain filling. Two ‘super’ rice cultivars were grown under two irrigation regimes of well-watered(WW) and alternate wetting and moderate soil drying(AWMD). Compared with the WW treatment,the activities of ADP glucose pyrophosphorylase(AGPase), starch synthase(StSase) and starch branching enzyme(SBE), and the accumulation of non-structural carbohydrates(NSCs) in the stems before heading were significantly improved, and more starch granules were stored in the stems in the AWMD treatment. After heading, the activities of α-amylase, β-amylase, sucrose phosphate synthase(SPS) and sucrose synthase in the synthetic direction(SSs)were increased in the stems to promote the remobilization of NSCs for grain filling under AWMD. During grain filling, the enzymatic activities of sucrose synthase in the cleavage direction(SSc), AGPase, StSase and SBE in the inferior spikelets were increased, which promoted grain filling, especially for the inferior spikelets under AWMD.However, there were no significant differences in vascular microstructures. The grain yield and grain weight could be improved by 13.1 and 7.5%, respectively, by optimizing of the irrigation regime. We concluded that the low activities of key enzymes in carbon metabolism is the key limitation for the poor grain filling, as opposed to the vascular microstructures, and AWMD can increase the amount of NSC accumulation in the stems before heading, improve the utilization rate of NSCs after heading, and increase the grain filling, especially in the inferior spikelets, by altering the activities of key enzymes in carbon metabolism.展开更多
Integrative cultivation practices(ICPs)are essential for enhancing cereal yield and resource use efficiency.However,the effects of ICP on the rhizosphere environment and roots of paddy rice are still poorly understood...Integrative cultivation practices(ICPs)are essential for enhancing cereal yield and resource use efficiency.However,the effects of ICP on the rhizosphere environment and roots of paddy rice are still poorly understood.In this study,four rice varieties were produced in the field.Each variety was treated with six different cultivation techniques,including zero nitrogen application(0 N),local farmers’practice(LFP),nitrogen reduction(NR),and three progressive ICP techniques comprised of enhanced fertilizer N practice and increased plant density(ICP1),a treatment similar to ICP1 but with alternate wetting and moderate drying instead of continuous flooding(ICP2),and the same practices as ICP2 with the application of organic fertilizer(ICP3).The ICPs had greater grain production and nitrogen use efficiency than the other three methods.Root length,dry weight,root diameter,activity of root oxidation,root bleeding rate,zeatin and zeatin riboside compositions,and total organic acids in root exudates were elevated with the introduction of the successive cultivation practices.ICPs enhanced nitrate nitrogen,the activities of urease and invertase,and the diversity of microbes(bacteria)in rhizosphere and non-rhizosphere soil,while reducing the ammonium nitrogen content.The nutrient contents(ammonium nitrogen,total nitrogen,total potassium,total phosphorus,nitrate,and available phosphorus)and urease activity in rhizosphere soil were reduced in all treatments in comparison with the non-rhizosphere soil,but the invertase activity and bacterial diversity were greater.The main root morphology and physiology,and the ammonium nitrogen contents in rhizosphere soil at the primary stages were closely correlated with grain yield and internal nitrogen use efficiency.These findings suggest that the coordinated enhancement of the root system and the environment of the rhizosphere under integrative cultivation approaches may lead to higher rice production.展开更多
The Pouni area is made up of basalts belonging to the Boromo belt, lamprophyres and granitoids. These geological formations are similar to geological formations of the same type in other regions of the Palaeoproterozo...The Pouni area is made up of basalts belonging to the Boromo belt, lamprophyres and granitoids. These geological formations are similar to geological formations of the same type in other regions of the Palaeoproterozoic domain of the Man/Leo shield. This study, which focused on the petrographic and geochemical characteristics of these geological formations, led to the following main conclusions: The lamprophyres are basic plutonic rocks that cut through other geological formations. The basalt belongs to the northern part of the Borormo belt and is thought to be a relic of overthickened oceanic plateaus. There are two groups of granitoid rocks. The granodiorite has a geochemical signature close to that of Archean TTGs and is metaluminous in character. It has a low potassium content. The minor element and rare earth element spectra indicate that it could be derived from partial melting of basic magmatic rocks. Biotite granites are peraluminous and highly potassic. Minor element contents and rare earth spectra indicate that they could be derived from partial melting of felsic materials. Geotectonic diagrams show that the granitoids identified in the Pouni zone were emplaced in an active tectonic context, similar to that of present-day subduction zones.展开更多
Alfalfa is the most widely cultivated perennial legume forage crop worldwide.Drought is one of the major environmental factors influencing alfalfa productivity.However,the molecular mechanisms underlying alfalfa respo...Alfalfa is the most widely cultivated perennial legume forage crop worldwide.Drought is one of the major environmental factors influencing alfalfa productivity.However,the molecular mechanisms underlying alfalfa responses to drought stress are still largely unknown.This study identified a drought-inducible gene of unknown function,designated as Medicago sativa DROUGHT-INDUCED UNKNOWN PROTEIN 1(MsDIUP1).MsDIUP1 was localized to the nucleus,chloroplast,and plasma membranes.Overexpression of MsDIUP1 in Arabidopsis resulted in increased tolerance to drought,with higher seed germination,root length,fresh weight,and survival rate than in wild-type(WT)plants.Consistently,analysis of MsDIUP1 over-expression(OE)alfalfa plants revealed that MsDIUP1 also increased tolerance to drought stress,accompanied by physiological changes including reduced malondialdehyde(MDA)content and increased osmoprotectants accumulation(free proline and soluble sugar),relative to the WT.In contrast,disruption of MsDIUP1 expression by RNA interference(RNAi)in alfalfa resulted in a droughthypersensitive phenotype,with a lower chlorophyll content,higher MDA content,and less osmoprotectants accumulation than that of the WT.Transcript profiling of alfalfa WT,OE,and RNAi plants during drought stress showed differential responses for genes involved in stress signaling,antioxidant defense,and osmotic adjustment.Taken together,these results reveal a positive role for MsDIUP1 in regulating drought tolerance.展开更多
Monogenic lines,which carried 23 genes for blast resistance were tested and used donors to transfer resistance genes by crossing method.The results under blast nursery revealed that 9 genes from 23 genes were suscepti...Monogenic lines,which carried 23 genes for blast resistance were tested and used donors to transfer resistance genes by crossing method.The results under blast nursery revealed that 9 genes from 23 genes were susceptible to highly susceptible under the three locations(Sakha,Gemmeza,and Zarzoura in Egypt);Pia,Pik,Pik-p,Piz-t,Pita,Pi b,Pi,Pi 19 and Pi 20.While,the genes Pii,Pik-s,Pik-h,Pi z,Piz-5,Pi sh,Pi 3,Pi 1,Pi 5,Pi 7,Pi 9,Pi 12,Pikm and Pita-2 were highly resistant at the same locations.Clustering analysis confirmed the results,which divided into two groups;the first one included all the susceptible genes,while the second one included the resistance genes.In the greenhouse test,the reaction pattern of five races produced 100%resistance under artificial inoculation with eight genes showing complete resistance to all isolates.The completely resistant genes:Pii,Pik-s,Piz,Piz-5(=bi2)(t),Pita(=Pi4)(t),Pita,Pi b and Pi1 as well as clustering analysis confirmed the results.In the F1 crosses,the results showed all the 25 crosses were resistant for leaf blast disease under field conditions.While,the results in F2 population showed seven crosses with segregation ratio of 15(R):1(S),two cross gave segregated ratio of 3 R:1 S and one gave 13:3.For the identification of blast resistance genes in the parental lines,the marker K3959,linked to Pik-s gene and the variety IRBLKS-F5 carry this gene,which was from the monogenic line.The results showed that four genotypes;Sakha 105,Sakha 103,Sakha 106 and IRBLKS-F5 were carrying Pik-s gene,while was absent in the Sakha 101,Sakha 104,IRBL5-M,IRBL9-W,IRBLTACP1 and IRBL9-W(R)genotypes.As for Pi 5 gene,the results showed that it was present in Sakha 103 and Sakha 104 varieties and absent in the rest of the genotypes.In addition,Pita-Pita-2 gene was found in the three Egyptian genotypes(Sakha 105,Sakha 101 and Sakha 104)plus IRBLTACP1 monogenetic.In F2 generation,six populations were used to study the inheritance of blast resistance and specific primers to confirm the ratio and identify the resistance genes.However,the ratios in molecular markers were the same of the ratio under field evaluation in the most population studies.These findings would facilitate in breeding programs for gene pyramiding and gene accumulation to produce durable resistance for blast using those genotypes.展开更多
Most indigenous rice landraces are sensitive to photoperiod during short day seasons,and this sensitivity is more pronounced in indica than in japonica landraces.Attempts to identify photoperiod sensitive(PPS)cultivar...Most indigenous rice landraces are sensitive to photoperiod during short day seasons,and this sensitivity is more pronounced in indica than in japonica landraces.Attempts to identify photoperiod sensitive(PPS)cultivars based on the life history stages of the rice plant,and several models and indices based on phenology and day length have not been precise,and in some cases yield counterfactual inferences.Following the empirical method of traditional Asian rice farmers,the author has developed a robust index,based on the sowing and flowering dates of a large number of landraces grown in different seasons from 2020 to 2023,to contradistinguish PPS from photoperiod insensitive cultivars.Unlike other indices and models of photoperiod sensitivity,the index does not require the presumed duration of different life history stages of the rice plant but relies only on the flowering dates and the number of days till flowering of a rice cultivar sown on different dates to consistently identify photoperiod sensitive cultivars.展开更多
基于开放式臭氧浓度升高O_3-FACE(Free-Air Concentration Elevation of O_3)实验平台,利用前期水稻O_3-FACE试验的基础数据,通过建立水稻产量与不同评价指标(累积气孔O_3吸收通量PODY和O_3浓度指标AOTX)的响应关系,比较了水稻产量损失...基于开放式臭氧浓度升高O_3-FACE(Free-Air Concentration Elevation of O_3)实验平台,利用前期水稻O_3-FACE试验的基础数据,通过建立水稻产量与不同评价指标(累积气孔O_3吸收通量PODY和O_3浓度指标AOTX)的响应关系,比较了水稻产量损失与各评价指标的相关性差异,通过对暴露剂量、吸收通量相关参数取值与产量损失的观察和分析结果的比较,找出更为合理的农作物臭氧风险评估阈值。结果表明:随着通量阈值Y[0~11 nmol O_3·m^(-2)PLA·s^(-1)(PLA:projected leaf area,投影叶面积)]和暴露浓度阈值X(0~50 n L·L^(-1))的增加,回归分析R^2值逐渐增加,当Y为11 nmol O_3m^(-2)PLA·s^(-1)和X为50 n L·L^(-1)时,气孔臭氧吸收通量POD11和累积暴露剂量AOT50与水稻相对产量的相关性最大,当通量阈值Y为8~13 nmol O_3·m^(-2)PLA·s^(-1)和暴露阈值X为46~58 n L·L^(-1)时,可获得较高的R^2值取值范围,分别为0.70~0.75和0.70~0.745。参考文献发现,目前地表臭氧污染可能引起的水稻产量损失范围为5%~8%,对照圈中POD9~10和AOT40~45产量损失的预测值亦在这区间,但前者R^2值(0.73~0.74)明显高于后者R^2值(0.64~0.69),表明基于气孔臭氧通量的评价指标能更好地反映水稻产量的变化。通过进一步分析发现,当通量阈值Y为9 nmol O_3·m^(-2)PLA·s^(-1)时,能更准确地评估水稻产量损失,且其R^2值(0.73)高于通量指标POD6(0.57)。以上研究结果表明,通量指标POD9更适合评估亚热带地区O_3污染对水稻作物的影响。展开更多
基金This work was supported by the National Key Research and Development Program of China(2022YFD1201600)the National Natural Science Foundation of China(32171964)the Science Fund for Creative Research Groups of Chongqing,China(cstc2021jcyj-cxttX0004)。
文摘Tillering is an important agronomic trait of rice(Oryza sativa)that affects the number of effective panicles,thereby affecting yields.The phytohormone auxin plays a key role in tillering.Here we identified the high tillering and semi-dwarf 1(htsd1)mutant with auxin-deficiency root characteristics,such as shortened lateral roots,reduced lateral root density,and enlarged root angles.htsd1 showed reduced sensitivity to auxin,but the external application of indole-3-acetic acid(IAA)inhibited its tillering.We identified the mutated gene in htsd1 as AUXIN1(OsAUX1,LOC_Os01g63770),which encodes an auxin influx transporter.The promoter sequence of OsAUX1 contains many SQUAMOSA PROMOTER BINDING PROTEIN-LIKE(SPL)binding sites,and we demonstrated that SPL7 binds to the OsAUX1 promoter.TEOSINTE BRANCHED1(OsTB1),a key gene that negatively regulates tillering,was significantly downregulated in htsd1.Tillering was enhanced in the OsTB1 knockout mutant,and the external application of IAA inhibited tiller elongation in this mutant.Overexpressing OsTB1 restored the multi-tiller phenotype of htsd1.These results suggest that SPL7 directly binds to the OsAUX1 promoter and regulates tillering in rice by altering OsTB1 expression to modulate auxin signaling.
基金This project was finically supported by the R&D Foundation of Jiangsu Province,China(BE2022425)the National Key Research and Development Program of China(2022YFD2300304)the Priority Academic Program Development of Jiangsu Higher-Education Institutions,China(PAPD).
文摘Recently developed ‘super’ rice cultivars with greater yield potentials often suffer from the problem of poor grain filling, especially in inferior spikelets. Here, we studied the activities of enzymes related to starch metabolism in rice stems and grains, and the microstructures related to carbohydrate accumulation and transportation to investigate the effects of different water regimes on grain filling. Two ‘super’ rice cultivars were grown under two irrigation regimes of well-watered(WW) and alternate wetting and moderate soil drying(AWMD). Compared with the WW treatment,the activities of ADP glucose pyrophosphorylase(AGPase), starch synthase(StSase) and starch branching enzyme(SBE), and the accumulation of non-structural carbohydrates(NSCs) in the stems before heading were significantly improved, and more starch granules were stored in the stems in the AWMD treatment. After heading, the activities of α-amylase, β-amylase, sucrose phosphate synthase(SPS) and sucrose synthase in the synthetic direction(SSs)were increased in the stems to promote the remobilization of NSCs for grain filling under AWMD. During grain filling, the enzymatic activities of sucrose synthase in the cleavage direction(SSc), AGPase, StSase and SBE in the inferior spikelets were increased, which promoted grain filling, especially for the inferior spikelets under AWMD.However, there were no significant differences in vascular microstructures. The grain yield and grain weight could be improved by 13.1 and 7.5%, respectively, by optimizing of the irrigation regime. We concluded that the low activities of key enzymes in carbon metabolism is the key limitation for the poor grain filling, as opposed to the vascular microstructures, and AWMD can increase the amount of NSC accumulation in the stems before heading, improve the utilization rate of NSCs after heading, and increase the grain filling, especially in the inferior spikelets, by altering the activities of key enzymes in carbon metabolism.
基金supported by the National Key Research and Development Program of China (2022YFD2300304)the National Natural Science Foundation of China (32071944 and 32272197)+2 种基金the Hong Kong Research Grants Council, China (GRF 14177617, 12103219, 12103220, and AoE/M-403/16)the State Key Laboratory of Agrobiotechnology (Strategic Collaborative Projects) in The Chinese University of Hong Kong, China, the Six Talent Peaks Project in Jiangsu Province, China (SWYY151)the Project Funded by the Priority Academic Program Development of Jiangsu Higher Education Institutions, China (PAPD).
文摘Integrative cultivation practices(ICPs)are essential for enhancing cereal yield and resource use efficiency.However,the effects of ICP on the rhizosphere environment and roots of paddy rice are still poorly understood.In this study,four rice varieties were produced in the field.Each variety was treated with six different cultivation techniques,including zero nitrogen application(0 N),local farmers’practice(LFP),nitrogen reduction(NR),and three progressive ICP techniques comprised of enhanced fertilizer N practice and increased plant density(ICP1),a treatment similar to ICP1 but with alternate wetting and moderate drying instead of continuous flooding(ICP2),and the same practices as ICP2 with the application of organic fertilizer(ICP3).The ICPs had greater grain production and nitrogen use efficiency than the other three methods.Root length,dry weight,root diameter,activity of root oxidation,root bleeding rate,zeatin and zeatin riboside compositions,and total organic acids in root exudates were elevated with the introduction of the successive cultivation practices.ICPs enhanced nitrate nitrogen,the activities of urease and invertase,and the diversity of microbes(bacteria)in rhizosphere and non-rhizosphere soil,while reducing the ammonium nitrogen content.The nutrient contents(ammonium nitrogen,total nitrogen,total potassium,total phosphorus,nitrate,and available phosphorus)and urease activity in rhizosphere soil were reduced in all treatments in comparison with the non-rhizosphere soil,but the invertase activity and bacterial diversity were greater.The main root morphology and physiology,and the ammonium nitrogen contents in rhizosphere soil at the primary stages were closely correlated with grain yield and internal nitrogen use efficiency.These findings suggest that the coordinated enhancement of the root system and the environment of the rhizosphere under integrative cultivation approaches may lead to higher rice production.
文摘The Pouni area is made up of basalts belonging to the Boromo belt, lamprophyres and granitoids. These geological formations are similar to geological formations of the same type in other regions of the Palaeoproterozoic domain of the Man/Leo shield. This study, which focused on the petrographic and geochemical characteristics of these geological formations, led to the following main conclusions: The lamprophyres are basic plutonic rocks that cut through other geological formations. The basalt belongs to the northern part of the Borormo belt and is thought to be a relic of overthickened oceanic plateaus. There are two groups of granitoid rocks. The granodiorite has a geochemical signature close to that of Archean TTGs and is metaluminous in character. It has a low potassium content. The minor element and rare earth element spectra indicate that it could be derived from partial melting of basic magmatic rocks. Biotite granites are peraluminous and highly potassic. Minor element contents and rare earth spectra indicate that they could be derived from partial melting of felsic materials. Geotectonic diagrams show that the granitoids identified in the Pouni zone were emplaced in an active tectonic context, similar to that of present-day subduction zones.
基金supported by the Strategic Pilot Projects of Chinese Academy of Sciences(XDA26030103)the National Natural Science Foundation of China(31722055 and 31672476)the Key Science and Technology Foundation of Gansu Province(19ZD2NA002)。
文摘Alfalfa is the most widely cultivated perennial legume forage crop worldwide.Drought is one of the major environmental factors influencing alfalfa productivity.However,the molecular mechanisms underlying alfalfa responses to drought stress are still largely unknown.This study identified a drought-inducible gene of unknown function,designated as Medicago sativa DROUGHT-INDUCED UNKNOWN PROTEIN 1(MsDIUP1).MsDIUP1 was localized to the nucleus,chloroplast,and plasma membranes.Overexpression of MsDIUP1 in Arabidopsis resulted in increased tolerance to drought,with higher seed germination,root length,fresh weight,and survival rate than in wild-type(WT)plants.Consistently,analysis of MsDIUP1 over-expression(OE)alfalfa plants revealed that MsDIUP1 also increased tolerance to drought stress,accompanied by physiological changes including reduced malondialdehyde(MDA)content and increased osmoprotectants accumulation(free proline and soluble sugar),relative to the WT.In contrast,disruption of MsDIUP1 expression by RNA interference(RNAi)in alfalfa resulted in a droughthypersensitive phenotype,with a lower chlorophyll content,higher MDA content,and less osmoprotectants accumulation than that of the WT.Transcript profiling of alfalfa WT,OE,and RNAi plants during drought stress showed differential responses for genes involved in stress signaling,antioxidant defense,and osmotic adjustment.Taken together,these results reveal a positive role for MsDIUP1 in regulating drought tolerance.
基金Authors extend their appreciation to Deanship of Scientific Research,King Faisal University,Saudi Arabia,for supporting this research(NA000112).
文摘Monogenic lines,which carried 23 genes for blast resistance were tested and used donors to transfer resistance genes by crossing method.The results under blast nursery revealed that 9 genes from 23 genes were susceptible to highly susceptible under the three locations(Sakha,Gemmeza,and Zarzoura in Egypt);Pia,Pik,Pik-p,Piz-t,Pita,Pi b,Pi,Pi 19 and Pi 20.While,the genes Pii,Pik-s,Pik-h,Pi z,Piz-5,Pi sh,Pi 3,Pi 1,Pi 5,Pi 7,Pi 9,Pi 12,Pikm and Pita-2 were highly resistant at the same locations.Clustering analysis confirmed the results,which divided into two groups;the first one included all the susceptible genes,while the second one included the resistance genes.In the greenhouse test,the reaction pattern of five races produced 100%resistance under artificial inoculation with eight genes showing complete resistance to all isolates.The completely resistant genes:Pii,Pik-s,Piz,Piz-5(=bi2)(t),Pita(=Pi4)(t),Pita,Pi b and Pi1 as well as clustering analysis confirmed the results.In the F1 crosses,the results showed all the 25 crosses were resistant for leaf blast disease under field conditions.While,the results in F2 population showed seven crosses with segregation ratio of 15(R):1(S),two cross gave segregated ratio of 3 R:1 S and one gave 13:3.For the identification of blast resistance genes in the parental lines,the marker K3959,linked to Pik-s gene and the variety IRBLKS-F5 carry this gene,which was from the monogenic line.The results showed that four genotypes;Sakha 105,Sakha 103,Sakha 106 and IRBLKS-F5 were carrying Pik-s gene,while was absent in the Sakha 101,Sakha 104,IRBL5-M,IRBL9-W,IRBLTACP1 and IRBL9-W(R)genotypes.As for Pi 5 gene,the results showed that it was present in Sakha 103 and Sakha 104 varieties and absent in the rest of the genotypes.In addition,Pita-Pita-2 gene was found in the three Egyptian genotypes(Sakha 105,Sakha 101 and Sakha 104)plus IRBLTACP1 monogenetic.In F2 generation,six populations were used to study the inheritance of blast resistance and specific primers to confirm the ratio and identify the resistance genes.However,the ratios in molecular markers were the same of the ratio under field evaluation in the most population studies.These findings would facilitate in breeding programs for gene pyramiding and gene accumulation to produce durable resistance for blast using those genotypes.
文摘Most indigenous rice landraces are sensitive to photoperiod during short day seasons,and this sensitivity is more pronounced in indica than in japonica landraces.Attempts to identify photoperiod sensitive(PPS)cultivars based on the life history stages of the rice plant,and several models and indices based on phenology and day length have not been precise,and in some cases yield counterfactual inferences.Following the empirical method of traditional Asian rice farmers,the author has developed a robust index,based on the sowing and flowering dates of a large number of landraces grown in different seasons from 2020 to 2023,to contradistinguish PPS from photoperiod insensitive cultivars.Unlike other indices and models of photoperiod sensitivity,the index does not require the presumed duration of different life history stages of the rice plant but relies only on the flowering dates and the number of days till flowering of a rice cultivar sown on different dates to consistently identify photoperiod sensitive cultivars.
