小麦发育过程及生育期机理模型的研究I.建模的基本设想与模型的描述

将小麦发育的温度效应曲线化,以发育生理生态过程为基础,利用作物生理发育时间(Physiological Development Time,简称PDT)为尺度,提出系统地预测小麦顶端发育阶段和物候生育期的模拟模型.预测的顶端发育阶段包括单棱期、二棱期、小花原基分化期、雌雄蕊原基分化期、药隔期、四分体期、抽穗期.物候发育阶段包括种子萌发、出苗期、分蘖期、越冬期、返青期、拔节期、孕穗期、抽穗期、开花期、灌浆期、成熟期.模型中用来描述特定品种发育遗传差异的参数有温度敏感性、生理春化时间、光周期敏感性和基本早熟性,分别体现了不同品种小麦在热效应、春化作用、光周期反应以及最早开花时间这四方面的遗传特性,共同决定了不同品种到达各发育阶段的生理发育时间     

基于生理生态过程的大麦顶端发育和物候期模拟模型检验

为测试研究I 模型BarleyGrow,采用4个生态区(南京、扬州、武汉、昆明)、10个大麦品种在不同播期下的顶端发育和物候期资料,对BarleyGrow、YDmodel和SUCROS模型进行对比检验和评价.利用遗传-模拟退火算法确定各品种的遗传参数,提高了应用程序求算参数的精度.从模型的整体预测效果来看,BarleyGrow对不同地区、不同播期、不同品种的各顶端发育和物候期预测准确而稳定,均方差RMSE在1.06～7.94d之间,而YDmodel为6.26～13.35d,SUCROS为11.22～20.28d.各参试品种对BarleyGrow中灌浆期基点温度、生理春化时间、临界日长、最短苗穗期4参数反应敏感.经改进的生理发育时间(PDT)模拟模型(BarleyGrow)对中国广大地区不同温光条件下的大麦顶端发育和物候发育均具有较好的预测效果,尤其对药隔期、二裂期、毛状期、抽穗期、灌浆期、成熟期的模拟精度高而稳定,表现出较强的机理性以及较好的预测性.     

小麦不同品种和播期对发育阶段的效应

以热时间（thermal time)为尺度研究了小麦不同品种和播期对发育阶段的效应,结果表明,小麦分蘖发生的早晚以生态因子调控为主,基因型差异较小;分薛- 节期为冬性品种（京411）一生中可变性最大的生育阶段,穗分化进入单棱斯的早晚以基因型效应为主,生态因子的影响次之,单棱－二棱期为春化作用的敏感期,冬性品种晚播（3月2日）春化效应可延迟到小花原基分化期之前,小麦物候期与穗发育阶段的对应关系具有一定的可变性,冬性品种较强的春化作用增加了其生态可变叶原基数;春化过程结束前,物候发育及穗发育阶段累计GDD与相应生殖器官原基分化的数的相关性不明显,春性品种（扬麦158）的物候发育及药隔分化期之前的穗发育阶段与各类顶端原基的分化数均具有极显著的正相关关系。     

大麦产量构成模型

通过产量构成法构建了适用于不同地区不同品种的大麦产量模拟模型.以南京、昆明、武汉3个地区各试验处理中不同大麦品种最适条件下的产量因素为基础,建立了最适条件下每株穗数相对值、每穗粒数相对值、千粒重相对值与累积光合有效辐射的回归方程,构建了实际条件下的不同大麦品种每株穗数、每穗粒数和千粒重与这三者在最适条件下的潜在值和实际条件下的水肥丰缺因子等变量的函数关系.模型较为全面地考虑了大麦品种生长发育的内外因素,内因主要体现为品种遗传特性（潜在的每株穗数、潜在的每穗粒数、潜在的千粒重和灌浆期因子）,外因包括光合有效辐射和水肥丰缺因子.运用武汉、昆明、扬州3个地区不同品种、不同播期的田间试验资料对模型进行了测试和检验.结果表明,模型对大麦产量构成因子及理论产量的模拟效果较好,模拟值与观测值吻合度高,每株穗数、每穗粒数和千粒重相对误差（RE）绝对值的平均值分别为1.96%、1.88%和1.67%,对这三者及理论产量的模拟值和观测值进行y=x回归分析,相关系数（r）在0.9464～0.9987,相关性显著,说明模型具有较高的预测性和适用性.     

大麦发育模型及其光敏感参数

为了确定光周期对作物影响的连续变化规律,采用大麦自交系为材料,通过大麦自交系从长日照到短日照和从短日照到长日照环境有规律地进行相互转移的试验方法,定量地研究了光周期对大麦自交系诱导的响应。结果表明:光周期对大麦生长发育的影响呈阶段性线性关系。据此将大麦花前期发育阶段划分为基本营养生长期(BVP)、光敏感期(PSP)和光敏感后期(PPP)。建立了光周期影响大麦发育的数学模型,计算出花前期各发育阶段的天数,然后根据大麦光温发育模拟模型确定模型参数:它们分别是f0(最短花前期)、θ1和θ2(光敏感期起始时间和光敏感期终止时间)、δ(光敏感系数);估出96个大麦品系(包括两个亲本)的发育模型参数。计算结果表明,96个大麦品系均受光周期的影响,不同的大麦品系感光性差异较大,最早的在出苗后9.78d就进入光敏感期,最迟的在播种后41.49d进入光敏感期;最快的9.2d通过光敏感期,最慢的光敏感期一直持续到抽穗前;不同的发育阶段光周期对大麦的影响也有明显的区别,表现为光敏感期强烈地受光周期的影响,而在基本营养生长期和光敏感后期的影响较小;并分析了大麦抽穗前各发育阶段的长短与感光性强弱的关系。     

基于生理发育时间和生长度日的烤烟生育期预测模型

利用2010-2011年在云南具有代表性的植烟县开展烤烟种植试验数据,建立基于生理发育时间和生长度日的生育期模型,再利用农业气象观测资料进行模型验证.结果表明：在移栽期之前,两种预测模型都有很好的模拟效果,平均误差日数较低,移栽期之后,受移栽、打顶等农事活动的影响,误差日数增大;使用生理发育时间法进行模拟的预测值与实测值符合度较好,尤其是在移栽期之前,平均误差日仅为2 d,预测精度明显高于生长度日法;受光周期效应的影响,生理发育时间模型在较低纬度地区的模拟效果偏差,而在较高纬度地区模拟精度较高.     

三种耧斗菜属植物柱头的特征

为了深入了解植物捕获花粉的机制和生殖策略,我们对3种耧斗菜属Aquilegia L.植物华北耧斗菜A. yabeana Kitagawa、无距耧斗菜A. ecalcarata Maxim.和秦岭耧斗菜A. incurvata P. K. Hsiao的雌蕊进行了扫描电镜和荧光显微镜观察,发现其柱头发育的特殊式样。其柱头位于花柱的腹缝面,由花柱腹缝两侧细胞发育成柱头乳突,在花期中从顶端朝花柱基部渐次成熟。伴随花柱向外弯曲,成熟的柱头表面相继呈现,逐渐增加了花粉落置的空间。这种柱头发育的式样通过延长柱头的可授期,增加了植物在不可预测的传粉环境下的繁殖机会。     

大麦叶面积指数模拟模型

准确模拟叶面积指数是作物生长模拟模型预测作物生长和产量的关键.本文通过系统分析扬州和武汉地区不同大麦品种高产群体叶面积指数变化动态,建立了大麦群体的叶面积指数模拟模型.大麦叶面积指数是品种叶面积指数扩展的遗传参数和气温日较差、日照时数、辐射量等气候因子及水肥丰缺因子的函数.孕穗抽穗期最大叶面积指数与该期最适叶面积指数是不同的概念,二者之间存在着极显著差异.利用扬州、南京和昆明地区不同品种的播期试验及氮肥试验资料对模型进行了检验,结果表明,模型对大麦叶面积指数的模拟效果较好,模拟值与观测值吻合度高,根均方差RMSE介于0.742～2.865,平均值为1.348.对模拟值与观测值进行y=x的线性回归分析,相关系数R²介于0.511～0.954,均呈极显著正相关.     

棉花生育时期及蕾铃发生发育模拟模型研究

通过定量分析南京、安阳、保定和石河子4个试验点2002年不同播期3个品种(早熟品种中棉所36号、中早熟品种中棉所35号、中熟品种中棉所41号)的生育时期与环境因子之间的动态关系,建立了基于生理发育时间(DPT)的棉花生育期、果枝出现时间及其蕾铃发育阶段的模拟模型.模型的热效应计算考虑了不同棉区昼夜温较差对棉花发育速率的影响以及薄膜覆盖的增温效应,在模型中引入了果枝始节系数(I_FIN)、日照时数因子(F_SH)和果枝节位光照系数(I_FBR).利用不同年份、生态区、基因型的试验资料对模型进行了测试检验.结果表明,不同条件下模拟值与观测值的符合度较好.不同生育时期的模拟值与观测值的根均方差(RMSE)从播种到出苗、出苗到现蕾、开花到吐絮及播种到吐絮分别为0.9、2.2、17和2.1d,平均2.1d;棉株各果节点从现蕾到开花日期的模拟值与观测值的RMSE为1.8～3.7d,从开花到吐絮日期的RMSE值为4.6～5.8d.     

超早熟栽培草莓花芽分化进程的扫描电镜观察

于2011年7月23日至8月24日,对草莓品种‘红颜’进行低温(夜温为17～20℃变温)、短日照(黑暗时间为16 h)处理,从处理之始到花序现蕾,约每3～5 d取样1次,实体解剖镜下解离草莓顶芽,材料均用FAA固定,临界点干燥,扫描电镜观察,以确定保护地栽培草莓的花芽开始分化始期,为生产中草莓尽早适期定植提供依据.根据扫描观察的微形态特征,将草莓花芽分化的全过程分为花芽未分化期、花芽分化始期、生殖顶端膨大期、花序分化期、顶花花萼与花瓣形成期、雄蕊形成期和雌蕊形成期共7个时期.花芽未分化期,顶芽一直处于营养生长阶段;花芽分化始期,草莓顶芽逐步由尖锐、狭窄的营养顶端向平坦、宽大的生殖顶端发育;生殖顶端膨大期,生殖顶端隆起而膨大;花序分化期,顶花序逐步分化出顶花原基和侧花原基;顶花花萼、花瓣形成期,花萼(包括副萼)、花瓣原基相继分化完成;雄蕊形成期,两轮雄蕊原基相继分化形成;雌蕊形成期,大量雌蕊原基逐步隆起,心皮原基逐步卷合、完成花芽分化.研究认为,草莓的定植期以生殖顶端充分膨大期为宜,即低温短日照处理15～20 d后即可定植大田.     

Genetics of Heading Time in Wheat (TRITICUM AESTIVUM L.). II. the Inheritance of Vernalization Response

The inheritance of vernalization response was studied in crosses involving four spring wheats (Sonora 64 (S), Pitic 62 (P), Justin (J) and Thatcher (T)) and three winter wheats (Blackhull (B), Early Blackhull (E) and Extra Early Blackhull (EE)).—All winter cultivars were highly responsive to vernalization, and Pitic 62 was the only spring cultivar whose time to heading was significantly accelerated following cold treatments. When vernalized and grown under long days, spring and winter cultivars became comparable in their heading response, indicating that cold requirement is the major attribute differentiating the heading behavior of true spring and true winter wheats.—Inheritance of growth habit in the F₁ generation of a five-parent diallel cross showed dominance of the spring character in all spring x winter crosses. Depending on the cross, one or two duplicate major genes governing growth habit were detected in F₂, F₃ and backcross generations grown in the field under long days in the absence of vernalizing temperatures. In some spring x winter crosses most of the variation in heading time among spring segregates could be attributed to the effects of major genes conditioning growth habit. In other crosses the heading patterns appeared more complex, indicating that genes with smaller effects are also involved in the control of heading response under spring or summer environments.—Evidence was presented supporting the hypothesis that the cultivar Pitic 62 carries a different allele at one of the two major loci governing its spring habit. This allele was associated with some response to vernalization and acted as a dominant gene determining earliness under low temperature vernalization, but as a partially recessive gene determining lateness in the absence of vernalizing temperatures. Genotypes were assigned to five cultivars as follows: S, CC DD; P, CC D'D'; J, cc DD; B and EE, cc dd.—The presence of major and minor genes and of multiple alleles governing response to photoperiod and vernalization was discussed in relation to the genetic manipulation of the heading response and to breeding wheat cultivars with specific or broad adaptation.     

Heading date QTL in a spring × winter barley cross evaluated in Mediterranean environments

Heading date is a key trait for the adaptation of barley to Mediterranean environments. We studied the genetic control of flowering time under Northern Spanish (Mediterranean) conditions using a new population derived from the spring/winter cross Beka/Mogador. A set of 120 doubled haploid lines was evaluated in the field, and under controlled temperature and photoperiod conditions. Genotyping was carried out with 215 markers (RFLP, STS, RAPD, AFLP, SSR), including markers for vernalization candidate genes, HvBM5 (Vrn-H1), HvZCCT (Vrn-H2), and HvT SNP22 (Ppd-H1). Four major QTL, and the interactions between them, accounted for most of the variation in both field (71–92%) and greenhouse trials (55–86%). These were coincident with the location of the major genes for response to vernalization and short photoperiod (Ppd-H2 on chromosome 1H). A major QTL, near the centromere of chromosome 2H was the most important under autumn sowing conditions. Although it is detected under all conditions, its action seems not independent from environmental cues. An epistatic interaction involving the two vernalization genes was detected when the plants were grown without vernalization and under long photoperiod. The simultaneous presence of the winter Mogador allele at the two loci produced a marked delay in heading date, beyond a mere additive effect. This interaction, combined with the effect of the gene responsive to short photoperiod, Ppd-H2, was found responsible of the phenomenon known as short-day vernalization, present in some of the lines of the population. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Flowering Behaviour of Four Annual Grass Species in Relation to Temperature and Photoperiod

Populations of four co-habiting annual grass species Bromusmollis L. (Soft brome), Hordeum hystrix Roth (Mediterraneanbarley grass), Lolium rigidum Gaud. (Wimmera ryegrass) and Vulpiabromoides (L.) S. F. Gray (Squirrel-tail fescue) were examinedfor the presence and comparative levels of vernalization andphotoperiod response. This was evaluated as the number of daysfrom sowing to heading in both long (16 h) and short (normal,over-winter) photoperiods at two levels of temperature. Wide variation among the species in both vernalization and photoperiodicresponse was detected. L. rigidum possessed a high level ofvernalization response and was comparatively sensitive to photoperiodwhile V. bromoides possessed little or no vernalization responseand was comparatively insensitive to photoperiod. B. mollisand H. hystrix appeared to be intermediate between these twospecies for both responses. There were wide differences in timeto heading under long photoperiod (16 h) and high temperature(20 °C) of plants derived from seed of three of the speciesripened under non-vernalizing temperatures. This variation indicatesthe likely existence of genetic differences in vernalizationresponse between plants of these populations. The implications of these findings to the adaptability of thesespecies to the Australian environment have been outlined. Bromus mollis L., soft brome, Hordeum hystrix Roth, Mediterranean barley grass, Lolium rigidum Gaud, Wimmera ryegrass, Vulpia bromoides (L.) S. F. Gray, Squirrel-tail fescue, flowering, vernalization, photoperiod, temperature     

The expected effects of climate change on wheat development

Air temperature and the atmospheric concentrations of carbon dioxide are expected to rise. These two factor have a great potential to affect development, growth and yield of crops, including wheat. Rising air temperature may affect wheat development more than rising atmospheric CO₂ as there is not yet evidence that elevated CO₂ concentrations can directly induce changes in wheat development. In winter wheat, temperature has a complex effect on development due to its strong interaction with vernalization and photoperiod. In this paper, potential effects of rising temperature on the development of winter wheat from sowing to heading are considered in the light of this complex controlling mechanism. Data from a large series of field trials made in Romania is analysed at first and, subsequently, the IATA-Wheat Phenology model is used to calculate the impact of air warming on wheat development under different climate change scenarios. Data from the field trials showed very clearly the occurrence of a complex temperature/photoperiod/vernalization interaction for field sown crops and demostrated that the photoperiodic and vernalization responses have a key role in controlling the duration of the emergence-heading period. Temperature plays, instead, a central role in controlling seed germination and crop emergence as well as leaf inititiation and leaf appearance rate. The results of model analysis showed very well that the impact of an even or uneven distribution of warning effects may be very different. In the first case, the model predicted that the duration of the vegetative period was at least partly reduced in some years. In the second case, the model suggested that if warming will be more pronounced in winter than in spring, as predicted for some areas of the world by General Circulation Models, we may expect an increase in the duration of the vegetative phase of growth. On the contrary, in case of a spring warming but unchanged winter temperatures, we may expect a substantial decrease in the duration of the vegetative period.     

广东早稻高产的适宜气温指标

选取了广东典型生态区的曲江、高要、广州、汕头和湛江5个代表性试验点进行地理分期播种试验,各试验点均采用3个主栽品种、设置6个播期,观测早稻不同发育期、测定其产量及其产量构成因子。结合附近气象站同期观测资料,采用数理统计和变异系数等方法,分析平均气温对早稻产量及产量构成因子的影响,确定了广东早稻高产不同生育期的适宜气温指标。结果表明:全生育期平均气温为23～24℃,其中出苗-分蘖期为18～21℃、分蘖-孕穗期为21～25℃、孕穗-齐穗期为24～28℃、齐穗-成熟期为27～30℃。     

Genotype Differences in Photoperiodic Sensitivity and Vernalization Response in Wheat

The regression of final leaf number on leaf number at transferfrom an 8 h to an 18 h photoperiod was used to compare the photoperiodicresponse of eight locally-adapted lines of wheat. Comparisonsof such regressions in the vernalized and univernalized conditionsenabled detection of the presence or absence of a significantcultivar vernalization response and comparisons of differingresponses between cultivars. Prior vernalization generally did not significantly alter the‘rate’ of photoperiodic response, as the slope ofthe LNT/FLN regression, indicating a certain physiological independenceof vernalization and photoperiodic responses in wheat. Differences, both in photoperiodic and vernalization responseof the eight wheats studied have been discussed in terms ofadaptability and breeding for maturity alteration in wheat.Evidence has been produced for the possible existence of a thirdfactor influencing developmental processes associated with floweringin wheat.     

播期对春小麦生长发育及产量的影响

为了给陇中半干旱区春小麦高产栽培提供依据,2010年在甘肃定西进行了春小麦分期播种试验,并对不同播期条件下春小麦生长发育及产量形成进行了分析。结果表明:随着播期的推迟,春小麦播种-抽穗期日数减少、全生育期明显缩短;5月下旬之前,越早播种的春小麦LAI越大,5月下旬之后,播种愈晚春小麦LAI越大。早播春小麦LAI峰值靠前,晚播峰值滞后;6月下旬之前,播期早的春小麦叶绿素含量高于播期晚的,6月下旬之后播期愈早叶绿素含量下降愈快;不同播期春小麦群体生长率和净同化率在孕穗-抽穗期后差异显著,表现为3月18日播期最大,4月7日播期最小。各播期干物质累积在拔节期后表现为快速递增趋势。在拔节期前,早播处理的干物质积累速率较慢。随着播期的推迟,单株干物质最大积累速率出现时间提前,籽粒最大灌浆速率出现时间推迟,千粒重表现为先升后降;灌浆3个阶段各参数受播期影响比较显著;早播春小麦产量最高。