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1.
赵翔  汪延良  王亚静  王西丽  张骁 《作物学报》2008,34(11):1970-1976
研究了Ca2+ 对NaCl胁迫下蚕豆气孔运动及质膜K+通道的影响。结果表明,100 mmol L-1 NaCl可明显诱导气孔开放,该现象可被10 mmol L-1 CaCl2 显著抑制。为探讨盐胁迫下Ca2+对K+和Na+跨膜运输的调控机制,我们利用膜片钳技术记录全细胞K+ 电流发现,在100 mmol L-1 NaCl胁迫下,加入10 mmol L-1 CaCl2胞外处理,显著抑制质膜K+内向及外向通道电流,这种抑制可被1 mmol L-1 La3+ (Ca2+通道抑制剂)缓解。非盐胁迫下,10 mmol L-1 CaCl2 胞外处理也能显著抑制质膜内向K+通道,但明显激活其外向通道,加入1 mmol L-1 La3+并不能被缓解。用H2O2专一的荧光探针二氯荧光素二乙酸酯(H2DCF-DA)单细胞分析保卫细胞内H2O2含量变化显示,在100 mmol L-1 NaCl盐胁迫下,10 mmol L-1 CaCl2 处理明显诱导H2O2在保卫细胞中积累;100 mmol L-1 NaCl和10 mmol L-1 CaCl2单独处理并不能诱导H2O2积累。推测Ca2+在盐胁迫下可能先诱导H2O2在胞内积累,进而激活质膜Ca2+通道,迅速提高胞内Ca2+浓度以抑制Na+通过质膜K+通道跨膜内流,同时调节Na+外流,两种效应共同作用促使气孔关闭,减少盐胁迫下水分的过度散失。上述结果将为Ca2+调控作物抗盐机制研究提供新的思路。  相似文献   

2.
玉米籽粒赖氨酸含量的遗传及其与产量的关系分析   总被引:3,自引:0,他引:3  
用5个不同基础群体的15个自选系作母本, 5个不同优势群的测验系作父本, 采用NCII交配设计配成75个杂交组合, 经1年2点田间试验, 用近红外光谱仪测定了亲本及其杂交种F1和F2籽粒的赖氨酸含量, 并用三倍体种子胚乳-细胞质-母体效应模型对赖氨酸含量进行遗传分析。结果表明, 玉米籽粒赖氨酸含量除受种子、母体效应和细胞质3套遗传体系共同控制之外, 还不同程度受环境因素的影响, 遗传主效应方差VG (VG=VA+VD+VC+VAm+VDm)占总遗传方差(VG+VGE)的76.3%, 其中, 种子效应方差(VA+VD)、细胞质效应方差(VC)和母体效应方差(VAm+VDm)分别占24.6%、19.7%和55.7%;赖氨酸含量以母体遗传力为主(h2m=40.98%), 其次为种子遗传力(h2o=17.86%), 而细胞质遗传力较低(h2c=14.29%)。同时发现籽粒赖氨酸含量与产量之间不存在明显相关(r = –0.027)。因此, 在高赖氨酸育种中, 必须重视高值亲本尤其是母本的选择以及不同优势类群间的广泛组配, 从变异广泛的基础材料中对赖氨酸含量和籽粒产量同时进行选择和改良是可能的。  相似文献   

3.
玉米种子萌发过程中Na+、K+和Ca2+含量变化与耐盐性的关系   总被引:1,自引:0,他引:1  
玉米耐盐品种登海9号和盐敏感品种浚单18在含0、50、100、150和200 mmol L-1 NaCl的营养液中萌发生长, 采用等离子质谱分别测定其萌动种子种皮、胚、胚乳和幼苗根、根颈、叶中Na+、K+、Ca2+的含量。结果表明, 随着培养液中NaCl浓度的增加, 玉米体内Na+含量逐渐升高, 在幼苗中表现地下部(根和根颈)显著高于地上部(叶); 在萌动种子中, 胚中Na+积累量显著高于种皮和胚乳。根系积累Na+能力较强, 胚拒Na+能力较弱, 种皮具有一定的Na+累积能力。随NaCl浓度的增加, K+和Ca2+含量逐渐降低, 尤其是Ca2+含量急剧减少, 达38.4%~55.9%(登海9号)和65.6%~78.2%(浚单18)。玉米根、根颈、种皮的Na+积累能力、叶的拒Na+能力和幼苗选择吸收Ca2+的能力可能与品种耐盐性有关。  相似文献   

4.
不同基因型陆地棉亲本及其杂交后代的耐盐性差异   总被引:4,自引:1,他引:3  
 以Bt抗虫棉选系抗96(父本)、普通陆地棉选系陆58(母本)及其杂交后代(F1、F2)为材料,利用不同盐分含量砂培和土培试验比较研究它们苗期的耐盐相对指标和某些耐盐理化指标变化。结果表明,随土壤盐浓度升高,棉花父、母本及其后代叶片内的K+含量均降低,而Na+含量均明显增加,K+/Na+显著变小,丙二醛、脯氨酸、可溶性糖和类胡萝卜素含量均不同程度增加,而净光合速率均降低。各品系的过氧化物酶活性均以0.23%NaCl处理较高,0.46%NaCl处理次之,对照最低。耐盐理化和综合指标皆以陆58最高、抗96最低、F1和F2介于其间。表明耐盐性以陆58最强、抗96最弱,F1和F2介于父母本之间,杂交后代的耐盐性受其亲本的限制。  相似文献   

5.
铝胁迫对花生根尖线粒体膜生理特性的影响   总被引:2,自引:1,他引:1  
线粒体在植物生命活动中发挥重要作用,以花生为材料,研究了在铝胁迫条件下,花生根尖细胞线粒体膜生理变化。结果表明,通过根长试验、苏木精染色和根尖铝离子含量测定,筛选到耐铝品种LH11,铝敏感品种R1549。铝胁迫后,两个品种根尖线粒体MDA含量增加,R1549的MDA含量均高于LH11,在处理浓度是20 μmol L-1和100 μmol L-1时,两品种的MDA含量差异显著,但在400 μmol L-1时,差异不显著;两品种根尖线粒体Ca2+-ATP酶活性和Ca2+含量呈下降趋势,且随铝溶液浓度增加而加快,R1549的线粒体Ca2+含量下降较LH11快;随处理铝溶液浓度增加,线粒体光密度持续下降,MPT不断增大,ΔΨm明显降低,线粒体中Cyt c/a减少,R1549较LH11下降更明显。试验结果说明在较高铝浓度胁迫下,两品种线粒体透性转换孔开放,膜透性增加,跨线粒体膜Ca2+转运系统活性降低,使胞质Ca2+超载,细胞色素C释放到细胞质中,诱导根尖细胞发生程序性死亡,从而抑制根生长;在低铝浓度下,与铝敏感品种相比,耐铝品种吸收铝少,脂质过氧化水平低,线粒体膜Ca2+-ATPase活性、MPTP和ΔΨm调控能力强,不易发生PCD,从而表现出较强的耐铝能力。  相似文献   

6.
在纯净空气与H2O/ CO2污染空气来流对比试验结果基础上,采用数值计算方法和化学动力学方法,研究了H2O和CO2污染组分对煤油燃料超声速燃烧的影响,获得了试验手段难以得到的燃烧室流场参数和性能数据。完成了相应的煤油燃料超声速燃烧室二维数值计算,其中匹配了进口总温、总压、马赫数、氧气摩尔分数和工作当量油气比。将数值计算结果与相应试验测量值进行了对比分析,并结合燃烧室流场数据、性能参数分析了H2O和CO2污染的动力学影响、以及对燃烧室性能的影响。研究表明:(1)数值计算结果与实验测量值总体上吻合,两种手段均体现了纯净空气来流时不同煤油当量油气比的燃烧室性能,并反映了一致的“污染效应”影响趋势;(2) H2O污染、H2O+ CO2污染的存在降低了煤油燃料超声速燃烧室性能,体现在燃烧诱导压升、燃烧效率、流向冲量增量的下降,而且随着污染组分含量的增加,燃烧室性能下降越加显著。   相似文献   

7.
室外盆栽条件下盐胁迫对甜高粱光系统II活性的影响   总被引:3,自引:0,他引:3  
室外盆栽条件下, 设置2个NaCl浓度(100 mmol L–1和200 mmol L–1), 调查盐胁迫对甜高粱光合特性和光系统II (PSII)活性的影响。结果表明,叶片Na+离子含量与Na+/K+比随盐浓度增加和处理时间延长而增加。净光合速率(Pn)、光系统II开放反应中心天线转化效率(Fv¢/Fm¢)、光化学猝灭系数(qP)和光系统II实际光化学效率(ΦPSII)随盐浓度的增加而降低,非光化学猝灭(NPQ)随盐浓度增加而增加;100 mmol L–1处理组的Pn、Fv¢/Fm¢、qP和ΦPSII随处理时间延长有所恢复,但200 mmol L–1处理组无此现象。光系统II (PSII)最大光化学效率(Fv/Fm)在100 mmol L–1 NaCl处理时影响较小,但在200 mmol L–1 NaCl处理时明显下降。短期盐胁迫未影响荧光诱导动力学曲线,而200 mmol L–1 NaCl处理5 d后荧光诱导动力学曲线O-K和O-J相上升。进一步研究证明,PSII的失活速率在两个盐浓度下均无明显变化,而修复速率在200 mmol L–1盐浓度处理5 d后降低明显。因此,认为室外盆栽条件下盐胁迫造成甜高粱碳同化能力降低并改变PSII激发能分配;叶片Na+离子含量的大幅增加会导致PSII活性下降及光抑制,这与PSII失活速率无关,主要是失活PSII修复速率受抑制的结果。这对理解户外盐胁迫条件下C4作物的光抑制机制具有一定意义。  相似文献   

8.

为了通过吸收光谱快速测量SO2胁迫下水稻叶片超氧化物歧化酶活性,采用小区试验研究了SO2毒害环境下水稻叶片超氧化物歧化酶活性与吸收光谱变量的相关性。结果表明:在整个生育期,相关曲线存在2个的波峰点,位于520~550 nm和720~750 nm 2个波段之间。计算720~ 750 nm和520~550 nm波段光谱吸收率的平均值A1A2,通过A1A2构建比值光谱变量(A1/A2)、相加光谱变量(A1+A2)及对数相加变量(lgA1+lgA2),利用光谱变量反演水稻叶片超氧化物歧化酶活性。研究发现:在水稻整个生育期,对数相加光谱变量与超氧化物歧化酶活性的相关性显著提高。选取对数相加变量构建水稻叶片超氧化物歧化酶活性的模型。为进一步提高吸收光谱快速测量SO2毒害环境下水稻叶片超氧化物歧化酶活性提出了探索性研究。

  相似文献   

9.
 选择4个棕色棉品种(系)与5个白色棉品种进行不完全双列杂交,得正反交组合各20个,测定了吐絮期正反交组合的净光合速率、胞间CO2浓度、叶绿素含量,叶绿素a/b、PSII的最大光化学效率,分析杂交F1代光合参数的杂种优势表现与正反交之间的差异。结果表明:吐絮期杂种F1各项光合参数没有明显的正向中亲优势、正向超亲优势。正反交组合之间,除胞间CO2浓度存在显著差异外,其它光合参数差异不显著,胞间CO2浓度受到细胞质效应影响。正交组合F1叶绿素含量与胞间CO2浓度存在显著正相关,反交组合F1净光合速率与胞间CO2浓度存在极显著正相关。  相似文献   

10.
高盐碱胁迫下野生大豆(Glycine soja)体内离子积累的差异   总被引:2,自引:0,他引:2  
在总含量3%高盐碱原土盆栽条件下,对搜集于津唐盐碱地895份野生大豆植株进行耐盐碱性评价。通过测定203株死亡及收获植株茎、叶中Na+、Cl、K+、Ca2+、Mg2+的含量,分析高盐碱胁迫下野生大豆植株离子含量的分布及野生大豆植株死亡和成熟时体内离子积累程度,并探讨耐盐碱型野生大豆的耐性机制。结果表明,野生大豆植株在营养生长期间,盐碱胁迫致死植株茎、叶无机离子含量在不同存活时间组间并未达显著水平,Na+和Cl积累达到一定含量即出现死亡现象,致死植株茎中Na+和Cl离子范围分别为3.239~4.682和4.639~6.328,叶中分别为1.754~2.349和4.126~5.073;能够存活到成熟的耐盐碱型野生大豆植株茎叶Na+和Cl含量存在低中高3种类型。高耐型野生大豆茎、叶平均Na+和Cl含量显著低于低耐盐型,且茎中K+和叶中Ca2+和Mg2+含量较高。在高耐型野生大豆植株茎叶中也存在Na+和Cl离子含量高水平和低水平两种类型,推测野生大豆可能存在两种耐盐碱机制,其一为高耐受性,其二为低吸收性。  相似文献   

11.
燕麦对碱胁迫的阳离子响应机制   总被引:3,自引:0,他引:3  
以耐碱性燕麦品种Vao-9和碱敏感性品种白燕5号为试验材料,采用盆栽法,用25、50、75、100 mmol L-1碱浓度(Na2CO3和NaHCO3按摩尔比1∶1混合)进行短期(14 d)和长期(28 d)胁迫处理,观测两品种根、茎、叶中Na+、K+、Ca2+、Mg2+吸收及分配特点,并从离子平衡吸收与分配角度,探讨燕麦对碱胁迫的生理适应机制。胁迫处理14 d后,燕麦体内Na+增加,K+下降,Ca2+和Mg2+变化不大,且两品种间各器官中4种离子的分配比例差异不显著。胁迫处理28 d后,两品种各器官中Na+增幅较大,K+、Ca2+和Mg2+降幅较大。Vao-9植株体内Na+、Ca2+含量大于白燕5号,但K+、Mg2+含量与白燕5号无显著差异,但两品种间各器官中4种离子的分配特点不同;当胁迫浓度达到100 mmol L-1时,与白燕5号相比,Vao-9叶片中少分配5.9个百分点Na+,多分配13.5个百分点K+、28.9Ca2+、10.9Mg2+,茎中多分配5.4个百分点Na+,少分配9.8个百分点K+,根中少分配28.9个百分点Ca2+、10.9Mg2+,因而Vao-9叶片中Na+ /K+、Na+ /Ca2+、Na+ /Mg2+值较白燕5号低。可见,燕麦通过提高阳离子选择吸收及器官分配能力以适应碱胁迫。  相似文献   

12.
X. Wang    Y. Jacob    S. Mastrantuono    J. Bazzano    R. Voisin  D. Esmenjaud 《Plant Breeding》2004,123(1):79-83
The spectrum and inheritance of resistance to the root‐knot nematode Meloidogyne hapla was studied in the diploid species Rosa multiflora and R. indica. The host suitability of seven rose rootstocks, namely five R. multiflora (K2, K1, Floradale, CE63 and CE65) and two R. indica (CE35 and Maroc) clonal accessions, was evaluated using four geographic isolates of M. hapla. Plants grown under greenhouse conditions were tested at high and durable inoculum pressure of nematodes and rated for nematode infestation five months after inoculation on a 0–5 gall index. Different host suitabilities to M. hapla were demonstrated depending on the nematode isolates: in R. multiflora, the clone K2 had a resistant (R) response to all isolates; the clone K1 ranged from intermediate (I) (isolate ‘Canada’) to resistant (other isolates), Floradale was shown to be intermediate, whereas CE63 and CE65 were moderate to good hosts (H). In R. indica, both rootstocks were good to excellent hosts for the isolate Canada but resistant to all three other isolates, thus expressing an isolate‐specific resistance. A study on the genetics of resistance in R. multiflora to the M. hapla isolate Canada was then conducted using an incomplete diallel cross involving all previous clones except Floradale. A total of 120 hybrid individuals belonging to several progenies representing the cross combinations R × I, R × H, I × H and H × H were evaluated. Individuals of each progeny generally ranged into a monomodal distribution that suggests polygenic inheritance of resistance. In the family Rosaceae, the differences in the resistance genetics to the meiotic species M. hapla and to the mitotic species, M. arenaria, M. incognita and M. javanica are discussed in relation to the reproductive status of the nematodes.  相似文献   

13.
Quinoa is recently introduced to Pakistan as a salt‐tolerant crop of high nutritional value. Open field trials were conducted to evaluate its performance on normal and salinity/sodicity‐degraded lands at two locations of different salinity/sodicity levels, S1 (UAF Farm, Normal Soil), S2 (Paroka Farm UAF, saline sodic), S3 (SSRI Farm, normal) and S4 (SSRI Farm, saline sodic) during 2013–2014. Two genotypes (Q‐2 and Q‐7) were grown in lines and were allowed to grow till maturity under RCBD split‐plot arrangement. Maximum seed yield (3,062 kg/ha) was achieved by Q‐7 at normal field (S1) soil which was statistically similar with yield of same genotype obtained from salt‐affected field S2 (2,870 kg/ha). Furthermore, low yield was seen from both genotypes from both S3 and S4 as compared to S1 and S2. Q‐7 was best under all four conditions. Minimum yield was recorded from Q‐2 (1,587 kg/ha) at S4. Q‐7 had higher SOD, proline, phenolic and K+ contents, and lower Na+ content in leaves as compared to Q‐2. High levels of antioxidants and K+/Na+ of Q‐7 helped to withstand salt stress and might be the cause of higher yields under both normal and salt‐affected soils. Seed quality (mineral and protein) did not decrease considerably under salt‐affected soils even improved seed K+, Mg2+ and Mn2+.  相似文献   

14.
S. W. Bang    K. Sugihara    B. H. Jeung    R. Kaneko    E. Satake    Y. Kaneko    Y. Matsuzawa 《Plant Breeding》2007,126(1):101-103
Intergeneric crosses were made between Brassica oleracea and Moricandia arvensis utilizing embryo rescue. Six F1 hybrid plants were generated in the cross‐combination of B. oleracea × M. arvensis from 64 pods by the placenta‐embryo culture technique, whereas three plants were produced in the reciprocal cross from 40 pods by the ovary culture technique. The hybrid plants were ascertained to be amphihaploid with 2n = 23 chromosomes in mitosis and a meiotic chromosome association of (0–3)II + (17–23)I at metaphase I (M I). In the backcross with B. oleracea, some of these hybrids developed sesquidiploid BC1 plants with 2n = 32 chromosomes that predominantly exhibited a meiotic configuration of (9II + 14I) in pollen mother cells. The following backcross of BC2 plants to B. oleracea generated 48 BC3 progeny with somatic chromosomes from 2n = 19 to 2n = 41. The 2n = 19 plants showed a chromosomal association type of (9II + 1I) and a chromosomal distribution type of (91/2 + 91/2) or (9 + 10) at M I and M II, respectively. These facts might suggest that they were monosomic addition lines (MALs) of B. oleracea carrying a single chromosome of M. arvensis that could offer potential for future genetic and breeding research, together with other novel hybrid progeny developed in this intergeneric hybridization.  相似文献   

15.
Field experiments were conducted at the Directorate of Rice Research experimental farm, ICRISAT campus, Patancheru, Hyderabad, during 1998–2000 for five consecutive seasons (three wet and two dry seasons) with five treatments [T1 – 100 % straw incorporation; T2 – 50 % straw incorporation; T3 – 100 % straw + green manure (GM) incorporation; T4 – 100 % straw burning and T5 – 100 % straw removal (control)] along with the recommended dose of fertilizers to evaluate the effect of different crop residue management (CRM) practices on yield components and yield of rice in rice–rice cropping sequence. The ammonium N measured at active tillering was higher in 100 % straw‐added plots over 50 % straw addition and straw removal with maximum values in the straw + GM‐incorporated plots. Among the yield components, tillers, panicles and spikelets were influenced from the second season of residue incorporation with significant increase in 100 % straw‐added treatments. The increase in tiller and panicle number could be attributed to the increased NH4‐N in these treatments, which is evident from the significant correlation between tiller number and NH4‐N (r = 0.82**) and panicle number and NH4‐N (r = 0.87**). The influence of residue treatments on rice grain yield was observed from the third season onwards where incorporation of straw alone or in combination with GM and burning of straw significantly increased grain and straw yields. Grain yield showed significant positive correlation with the number of tillers (r = 0.74*–0.81**) and panicles (r = 0.74*–0.84**) in three treatments (T1, T3 andT4) where grain yields were significantly higher. The regression analysis showed that 57–66 % and 64–75 % of the variation in yield could be explained by tillers and panicles together in these three treatments during wet and dry seasons respectively. Thus, CRM practices such as addition of 100 % straw either alone or with GM and straw burning influenced the yield components (tillers, panicles and spikelets) positively and thereby increased rice grain yields.  相似文献   

16.
以苏玉糯4号为材料,采用二因素裂区设计,研究了不同基肥配比(纯N 75 kg hm-2、纯N 75 kg hm-2+K2O 70 kg hm-2、纯N 75 kg hm-2+P2O5 65 kg hm-2和纯N 75 kg hm-2 +P2O5 65 kg hm-2+K2O 70 kg hm-2)和拔节期追氮(0、150和300 kg hm-2)对糯玉米淀粉胶凝和回生特性的影响。结果表明,淀粉和回生淀粉的起始温度、峰值温度和终值温度虽然受到施肥处理的影响,但总体上变异较小。热焓值受基肥配比影响较小,糊化范围和峰值指数受拔节期追氮量影响较小。在不同基肥配比处理下,糊化范围在基施氮钾时最低,氮磷钾合理配施处理下最高,峰值指数表现和糊化范围相反。在拔节期不同追氮量处理下,热焓值以追氮150 kg hm-2时最高,追氮300 kg hm-2或不追氮无显著差异。胶凝淀粉冷藏后发生回生,表现为转变温度、热焓值和峰值指数降低,糊化范围变宽。和仅基施氮相比,增施磷或(和)钾都可降低淀粉的回生值和回生淀粉的热焓值,拔节期追氮处理的这两项指标均劣于不追氮处理。回生值与回生淀粉的热焓值和峰值指数显著正相关,相关系数分别为0.90 (P < 0.01)和0.41 (P < 0.05); 原淀粉的热焓值与峰值指数极显著正相关, 相关系数为0.65 (P < 0.01),与回生淀粉热焓值显著正相关,相关系数0.44 (P < 0.05),与终值温度显著负相关,相关系数-0.41 (P < 0.05)。在本试验条件下,以氮磷钾均衡基施并拔节期追氮150 kg hm-2时,淀粉胶凝和回生特性较为理想,表现为热焓值较高,回生值较低。  相似文献   

17.
以杂交棉石杂2号和新陆早43的F1、F2代及亲本NT2、H2、4-14为试材,通过测定不同生育时期各材料叶面积指数(LAI)、叶倾角(MTA)、冠层光截获率等指标,分析了各指标变化对群体光合物质生产的影响。结果表明,2个杂交棉F1代LAI具有超亲优势,冠层光截获率具有中亲优势; LAI和冠层光截获率具有明显的母系遗传特性,而MTA受到父本的显著影响。F2代冠层结构主要受F1代相关指标和衰退率的影响,LAI中亲优势减小了衰退率; 杂交棉F1代光合物质积累主要受亲本参数和超亲优势的影响,F2代主要受F1代参数的影响。杂交棉光合物质积累最大增长速率和直线增长期开始时间较晚,直线增长期及活跃增长期较长,最终积累量和最大增长速率较高。杂交棉F1代具有明显的光合生产和产量优势,F2代具有一定的产量优势。以选择具有优化冠层结构的亲本为基础,组配具有较大MTA的父本和较大LAI的母本,有利于改善杂交棉光合性能,提高群体光能利用率,进一步挖掘产量潜力,为杂交棉高光效组合的选育及提高F2代应用提供理论依据。  相似文献   

18.
Summary Interspecific substitutions of the nucleus of Helianthus annuus (2n=34) into the cytoplasm of H. petiolaris (2n=34) were obtained by successive backcrossing using cultivated sunflower, H. annuus, as the recurrent pollen parent.Meiosis in the F1 was characterized by multivalents, suggesting that 10 of the 34 chromosomes were heterozygous for chromosomal interchanges. An additional pair of chromosomes also contained a paracentric inversion. Continued backcrossing resulted in rapid elimination of the meiotic aberrations evident in the F1. In the BC1, 1 of 11 plants had normal meiosis and by the BC2, only 13 of 54 plants had meiotic aberrations similar to those of the F1. However, trisomic progeny (2n=35) were found in 3 of the 11 BC1 plants and 20 of the 54 BC2 plants. No meiotic aberrations were observed in BC3 or BC4 plants. Plants with indehiscent anthers, and considered to be male sterile (M.S.), first occurred in the BC1 and, by the BC2, 51 of 54 plants were M.S. All 19 BC3 and 16 BC4 plants were M.S. Preliminary investigations suggest that the pollen from such plants is sterile and that the sterility is cytoplasmic rather than genetic.Disc-flower measurements were a useful technique for selecting samples at the correct stage of microsporogenesis, but could not be used to distinguish between successive backcrosses.  相似文献   

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