小麦茎顶端原基分化的综合模式

研究了小麦 (TriticumaestivumL .)茎顶端不同类型原基分化的动态过程 ,以明确原基分化的综合模式 ,并建立了不同原基分化之间的定量关系。结果表明 ,小麦叶原基和苞叶原基分化与播后累积生长度日 (GDD ,growingdegreedaysaftersowing)的关系呈S形曲线 ,而小穗原基和小花原基为上升段抛物曲线。从分化模式看 ,苞叶原基具备营养器官原基特征 ;小穗和小花原基的分化进程能较好地反映基因型和生态条件对顶端发育的影响。小麦茎顶端原基分化的综合模式为由三段子模式构成的近似S曲线。叶原基数由基因型和环境条件共同决定 ,而苞叶原基、小穗原基和小花原基数以环境因子的影响为主。以平均热间距来衡量 ,适期播种处理的叶片、苞叶和小穗原基分化速率最高 ;而小花原基数与小花分化持续期之间的数量关系最为密切。研究结果有助于揭示和理解小麦茎顶端发育的生物学规律。     

Morphological Analysis of Leaf and Tiller Number Dynamics of Wheat (Triticum aestivumL.): Responses to Temperature and Light Intensity

In recent literature on Gramineae species, leaf and tiller numberdynamics have been studied by analysing site filling and thephyllochron of the mainstem. However, site filling is influencedby three components: (1) the phyllochron of the mainstem anddaughter tillers; (2) specific site usage (i.e. fraction ofbuds that ultimately develop into a visible tiller at a specificsite); and (3) HS-delay (i.e. difference in Haun Stage (HS)between the parent tiller and daughter tiller above the pointwhere the daughter tiller appears). These three morphologicalcomponents affecting site filling were studied under differentenvironmental conditions in a growth chamber experiment withspring and winter wheat (Triticum aestivumL.). Treatments weretemperature (daily average 10.5, 15.5 or 20.5 °C) and lightintensity (111, 191 or 286 µmol m^-2s^-1). Effects of temperatureand light intensity on phyllochron were well described by equationsalready reported in the literature. Specific site usage washigher at cooler temperatures and greater light intensitiesand was related to tiller position. It is proposed that theseeffects on specific site usage reflect differences in availabilityof local assimilate for tiller appearance. HS-delay of a tillerwas shorter if the expected tiller appearance was later andwas only slightly affected by light intensity or temperature.This new concept, combining HS-delay and specific site usage,can be useful in constructing more general models of the effectsof environmental factors on the dynamics of leaf number andleaf area ofGramineaespecies.Copyright 1998 Annals of BotanyCompany Triticum aestivum; wheat; phyllochron; temperature; light intensity; leaf number; tillering; site filling; site usage.     

Leaf Primordium Initiation and Expanded Leaf Production are Co-ordinated through Similar Response to Air Temperature in Pea (Pisum sativumL.)

Accurate prediction of the timing of leaf area development isessential to analyse and predict the responses of crops to theenvironment. In this paper, we analyse the two processes determiningthe chronology of leaf development—initiation of leafprimordia by the shoot meristem and production of expanded leavesout of the shoot tip—in several pea (Pisum sativumL.)cultivars in response to air temperature and plant growth rate.Contrasting levels of air temperature and plant growth rateduring leaf development were induced by a wide range of sowingdates and plant densities in glasshouse or field experiments.Full leaf expansion was found to occur one phyllochron afterfull leaf unfolding, whatever the leaf nodal position. Primordiuminitiation and expanded leaf production rates presented similarquantitative responses to air temperature (linear response andcommonx-intercept), whatever the plant growth rate, cultivaror period of cycle. As a consequence, they were co-ordinatedand the numbers of initiated primordia or expanded leaves wereeasily deduced from simple visual observation of leaf unfolding.The change, over time, of the numbers of initiated leaf primordiaand fully expanded leaves correlated with cumulated degree-days,with stable relationships in a wide range of environmental conditions.Two phases, with different production rates, had to be considered.These results allowed us to predict accurately the beginningand the end of individual leaf development from daily mean airtemperatures. The relationships obtained here provide an effectiveway of analysing and predicting leaf development responses tothe environment. Pisum sativumL.; pea; number of leaf primordia; number of leaves; temperature; modelling     

稻麦理论分蘖出叶数及其增长公式

稻麦作物分蘖出叶的数值化研究是作物形态发生光合功能等研究的一个重要基础．根据稻麦叶蘖同伸规律,设n为主茎总叶龄,Li为第i次分蘖上的理论出叶数,K为分蘖的最高次数,则Li和K均是n的函数．如果主茎芽鞘节和分蘖的芽鞘节均不发生分蘖,则K=（n-1）/3,L_i=C~i 1_n-2i(i=1,2,…,K）;如果主茎芽鞘节和分蘖鞘节均能发生分蘖,则K'=（n-1）／2,L'_i=C~i 1_n-i,以上K值均只取整数．分蘖与出叶增长的理论公式可用T＝∑sumfromi=0tok()C~i_A-2i（无鞘叶分蘖）或T'=sumfromi=0tok'()C_A-i-1~i,L'=sumfromi=0toK'()C_A-i-1~i,L'=sumfromi=0toK'()C_A-1~i 1（有鞘叶分蘖）来表示．这里T与L（T'与L'）分别为单株总茎蘖数与单株总叶数,A为叶龄．文章还采用数学归纳法对公式进行了数学证明,并展述了可能的应用前景．同样方法可证明莫惠栋等提出的稻麦理论分蘖数计算公式.     

Influence of Leaf Tolerance Mechanisms and Rain on Boron Toxicity in Barley and Wheat

Boron (B) toxicity is common in many areas of the world. Plant tolerance to high B varies widely and has previously been attributed to reduced uptake of B, most commonly as a result of B efflux from roots. In this study, it is shown that the expression of genes encoding B efflux transporters in leaves of wheat (Triticum aestivum) and barley (Hordeum vulgare) is associated with an ability of leaf tissues to withstand higher concentrations of B. In tolerant cultivars, necrosis in leaves occurred at B concentrations more than 2-fold higher than in sensitive cultivars. It is hypothesized that this leaf tolerance is achieved via redistribution of B by efflux transporters from sensitive symplastic compartments into the leaf apoplast. Measurements of B concentrations in leaf protoplasts, and of B released following infiltration of leaves, support this hypothesis. It was also shown that under B-toxic conditions, leaching of B from leaves by rain had a strong positive effect on growth of both roots and shoots. Measurements of rates of guttation and the concentration of B in guttation droplets indicated that the impact of guttation on the alleviation of B toxicity would be small.Boron (B) toxicity affects a wide variety of plants growing on soils with naturally high levels of B or when irrigated with water containing elevated levels of B (Stangoulis and Reid, 2002). Symptoms are most commonly seen as necrosis on leaf margins or leaf tips, depending on the type of leaf venation (Oertli and Kohl, 1961). Plant tolerance to high B varies considerably but is most commonly associated with reduced accumulation of B (Nable et al., 1997). Hayes and Reid (2004) identified differences in B efflux in roots as the primary determinant of the net uptake of B in barley (Hordeum vulgare). Reid (2007) established that this was also the mechanism for differences in B uptake in wheat (Triticum aestivum) and showed that there was a strong correlation between tolerance in both wheat and barley with the expression in roots of the genes TaBOR2 and HvBOR2, which encode B efflux transporters with homology to B efflux transporters in Arabidopsis (Arabidopsis thaliana) and rice (Oryza sativa; Takano et al., 2002; Nakagawa et al., 2007). Since the concentration of B in shoots was closely related to the concentration of B in roots (Hayes and Reid, 2004; Reid, 2007) a simple mechanism of tolerance could be explained. A similar mechanism of tolerance was shown to occur in Arabidopsis when roots overexpressed AtBor4 (Miwa et al., 2007).Sutton et al. (2007) made a qualitative analysis of the expression in leaves of Bot1 (which is identical to HvBOR2 and to avoid confusion will henceforth be referred to as HvBOR2) and found strong expression associated with hydathodes in the leaf tip. They proposed that in addition to root-based tolerance conferred by pumping of B from roots, that further tolerance could be achieved by excretion of B from hydathodes and its subsequent removal by rain. Oertli (1962) demonstrated that in young barley seedlings, significant amounts of B could be lost from leaves in this way.In the early work on B tolerance in cereals, it was noted that toxicity for plants grown in the field was generally observed at much lower concentrations of B in leaves than for plants grown in the glasshouse. For example, Nable et al. (1990) found that a 17% reduction in yield of field-grown barley occurred with a shoot B concentration of 62 mg kg⁻¹ dry weight (DW) whereas in the glasshouse the corresponding concentration was 120 mg kg⁻¹ DW. It was concluded that the most likely cause of the difference in shoot B between the growth conditions was leaching of B from leaves by rain in the field. However, an experiment in which a comparison was made between plants on which the leaves were regularly sprayed with water or not sprayed failed to show any difference in growth, despite significant reductions in leaf B in the sprayed plants (Nable et al., 1990).Jefferies et al. (1999) identified chromosome regions associated with tolerance in barley. They found a major locus on chromosome 4 that was related to reduced B uptake and a decrease in leaf symptoms. This locus was subsequently found to contain HvBOR2 (Sutton et al., 2007), whose expression in roots could explain both reduced B uptake and the decrease in leaf symptoms. In addition to the locus on chromosome 4, there was another locus on chromosome 2 that was associated with leaf symptom score but not associated with whole shoot B concentration (Jefferies et al., 1999).In this study we have shown that the expression of B efflux transporter genes in leaves results in enhanced tolerance to B, and contrary to previous reports, that rain can significantly reduce B toxicity.     

Initiation of Procambial Strands in Leaf Primordia of Bread Wheat, Triticum aestivum L

In Triticum aestivum L. the median and lateral procambial strandsserving the primordia originate independently and in isolationfrom the vascular system of the rest of the plant. The medianstrand is initiated first, followed by a succession of lateralstrands during the next four or so plastochrones. The medianand first lateral strands have their point of origin in theaxis, in the disc of insertion of the primordium. The laterlaterals are initiated up in the primordium. Once initiatedthe procambial strands extend from their point of origin bothacropetally and basipetally, the latter extension eventuallylinking them to strands associated with older leaves. It wouldappear that the materials necessary for the growth of the apicaldome and of the first four leaf primordia are supplied by generaldiffusion and not via direct vascular connexions with the restof the plant.     

Haustorium Formation and Cell Wall Appositions in Susceptible and Partially Resistant Wheat and Barley Seedlings Infected with Wheat Leaf Rust

Phase contrast light microscopy observations of wheat and barley seedlings infected with wheat leaf rust spores suggested that cell wall appositions are structural barriers against haustorium formation leading to abortion of infection structures. Nearly equal numbers of cell wall appositions per infection structure were detected in seedlings of susceptible and partially resistant wheat genotypes. Differences between susceptible and partially resistant genotypes became evident after the first haustorium had been formed. This again indicates the presence of a post-haustorial effect of partial resistance. Some factors influencing nutrient uptake are discussed. Wheat leaf rust colonies hardly formed haustoria in barley seedlings, the few not aborted infection structures were accompanied by cell collapse. The mechanisms of partial resistance in wheat and barley to their respective leaf rust fungi seem different, but their non-host reactions appear similar.     

Rate of Leaf Appearance and Final Number of Leaves in Wheat: Effects of Duration and Rate of Change of Photoperiod

This study was conducted to test the hypothesis that photoperiodor its rate of change significantly affects the rate of leafappearance (RLA) and final number of leaves (FNL) in wheat,as suggested from several time-of-sowing experiments. Two wheatcultivars (Condor and Thatcher) were sown in the field on 2Sep. 1992 at Melbourne (38°S). Photoperiod was extendedartificially to give five treatments up to terminal spikeletinitiation (TS) viz.: natural photoperiod (rate of change ofphotoperiod = 2 min d^-1), two faster rates of change (8·5and 13·3 min d^-1) and two constant photoperiods of 14·0and 15·5 h. After TS, the two constant photoperiods wereextended to 15·0 and 16·5 h, respectively, andtreatments were re-randomised, i.e. some plots received differentphotoperiod regimes before and after TS. The rate of leaf appearance maintained strong linear relationshipswith thermal time. It was greater for Condor [0·012-0·013(°C d)^-1] than for Thatcher [0·011-0·012 (°Cd)^-1] and did not alter during plant development or in responseto the change in photoperiod at TS. Rate of leaf appearanceon the main culm was not influenced by the rate of change ofphotoperiod nor by the average photoperiod. Cultivar and photoperiod significantly affected FNL on the mainculm. Condor produced more leaves than Thatcher under long butnot under short photoperiods. The rate of change of photoperioddid not affect FNL independently of the effect of average photoperiod.Most of the variation in FNL due to photoperiod resulted fromdifferences in duration of leaf initiation. The lack of effects of the photoperiod treatments on RLA contrastwith previous reports of its effects on the rate of phasic developmentfrom seedling emergence to double ridge. Therefore, the numberof visible leaves on the main culm (NL) at double ridge andat TS were not constant. However, NL on the main culm at doubleridge was closely correlated with FNL.Copyright 1994, 1999 AcademicPress Triticum aestivum L., wheat, leaf appearance, phyllochron, photoperiod     

CO2倍增对不同氮水平下小麦幼苗根系及叶片NR活性的影响

以小麦品种'小偃22'幼苗为材料,采用开顶式气室和水培实验研究了不同供氮水平(2.5、5.0、10.0和 15.0 mmol·L-1)下小麦幼苗植株生长量、根系形态、有机碳分泌速率和硝酸还原酶(NR)活性对大气CO2浓度升高的响应.结果显示,大气CO2浓度倍增均增加了小麦幼苗各生长阶段根冠生物量以及根系长度、面积、有机碳分泌速率和叶片NR活性.随供氮水平的提高,各生长阶段幼苗根冠生物量、根长和面积以及叶片NR活性呈上升趋势,而有机碳分泌速率呈下降趋势;根冠比变化不同阶段表现不一致,一叶一心期呈下降趋势,二叶一心期和三叶一心期分别以15.0和10.0 mmol·L-1氮水平较高.研究表明,大气CO2浓度升高可促进小麦幼苗根系生长和有机碳分泌速率,提高其氮素同化能力;增加介质供氮有利于高CO2浓度条件下小麦幼苗根冠生长和氮素同化,提高根冠比,减少根系有机碳过度分泌引起的碳损耗.     

Growth of Tiller Buds in Barley: Effects of Shade Treatment and Mineral Nutrition

Examination of the stem apex of Proctor barley showed that thebud of the coleoptile tiller, Tc, is probably present in thedry grain and that the bud, TI, carried in the axil of the firstleaf is present at or soon after 24 h from planting. Subsequentlytiller buds are initiated with a plastochron of about 4 days,this being rather longer than that for the foliar primordia.During the initial phase of bud growth vascular connectionsare established with the leaf above, but not to the subtendingleaf. At some time after these vascular connections are formedand when it has a dry weight of 4–7 µg the bud entersa phase of rapid, exponential growth in dry weight. Shading the first leaf delays the onset of rapid growth forboth Tc and Ti, but after a lag period rapid growth commences;this is coincident with development of the second leaf as anorgan exporting assimilated carbon. The phase of rapid growth of tiller buds is delayed when applicationof either nitrogenous or nonnitrogenous minerals is delayed.Ammonium was found to be less satisfactory as a nitrogen sourcethan nitrate, probably because of toxicity effects. Slight growthof Tc and T1 occurs in presence of nonnitrogenous minerals andabsence of nitrogen but growth is greater when nitrogen is suppliedin absence of the other minerals, although such growth is substantiallyless than that found when all nutrients are supplied. The interactionbetween nitrogen and non-nitrogenous minerals which controlsbud growth was not found to affect growth of the parent plantwhich is, as previously shown, controlled by timing of the nitrogensupply. AnotheT distinction is that higher concentrations ofnitrogen and the other minerals are required for maximum growthof the bud than for that of the plant. Tiller bud growth is interpreted as occurring in two phases.In the first, initiation, phase there is a close associationwith the subtending leaf, and nutritionally bud and leaf arelinked. This phase is followed by one in which the bud is directlyconnected by vascular traces to the leaf above, which becauseof this controls bud growth by modulating supply of assimilatedcarbon and nitrogen, and other minerals to it.     

Growth-Regulating Substances and the Growth of Tiller Buds in Barley; Effects of Cytokinins

An hypothesis was set up from which it was predicted that applicationof cytokinin to barley seedlings grown without mineral nutrientswould lead to rapid growth of the coleoptile and first leaftiller buds. Application of cytokinins to the leaves was ineffective,but supplying a number of known cytokinins by steeping the rootsof 4 d old seedlings in solution for 4 h led to significantgrowth of the coleoptile bud. Adenine and cytokinin analogueshad no effect. Supplying cytokinins through the roots also furtherenhanced the growth of buds of plants given mineral nutrients.Cytokinin treatment reduced root dry matter, with small reductionsin mean axis length and number of lateral roots. For plantsnot given mineral nutrients reduction in root weight was compensatedby an increase in weight of the aerial parts; however, for plantssupplied with mineral nutrients this was not so and the lowerroot weight resulted in a smaller total plant dry weight. An interpretation of tiller bud growth in terms of control byinteracting effects of mineral nutrition, assimilate supply,and cytokinin availability is proposed.     

Epidermal Cell Division and the Coordination of Leaf and Tiller Development

Initiation and development of grass leaves and tillers are oftendescribed individually with little attention to possible interrelationshipsamong organs. In order to better understand these interrelationships,this research examined epidermal cell division during developmentaltransitions at the apical meristem of tall fescue (Festuca arundinaceaSchreb.). Ten seedlings were harvested each day for a 9-d period,and lengths of main shoot leaves and primary tillers were measured.In addition, numbers and lengths of epidermal cells were determinedfor 0·5 mm segments along the basal 3 mm of each leafand tiller. Primordia development and onset of rapid leaf elongationwere characterized by an increase in the number of cells perepidermal file with mean cell length remaining near 20 µmper cell. After the leaf had lengthened to 1-1·5 mm,cells near the leaf tip ceased dividing and increased in length,at which time leaf elongation rate increased rapidly. Liguleformation, marking the boundary between blade and sheath cells,occurred prior to leaf tip emergence above the whorl of oldersheaths, while the earliest differentiation between blade andsheath cells probably began when leaves were < 1 mm long.Major transitions in leaf and tiller development appeared tobe synchronized among at least three adjacent nodes. At theoldest node, cessation of cell division in the leaf sheath wasaccompanied by initiation of cell division and elongation inthe associated tiller bud. At the next younger node the ligulewas being initiated, while at the youngest node cell divisioncommenced in the leaf primordium, as elongation of a new leafblade began. This synchronization of events suggests a key rolefor the cell division process in regulating leaf and tillerdevelopment.Copyright 1994, 1999 Academic Press Festuca arundinacea Schreb., tall fescue, cell division, leaf initiation, tillering, ligule development     

Tillering Potential and Relationship Between Leaf and Tiller Production in Perennial Ryegrass

A detailed morphological examination of the tillering characteristicsof perennial ryegrass (Loltum perenne L.) is presented. It isshown that the tillering potential of perennial ryegrass interms of site filling is 0–693 tillers/tiller/leaf appearanceinterval. This is higher than earlier presented values and isdue to the tillering ability of the prophyll bud which has notpreviously been taken into account. In a controlled climateroom experiment with spaced plants of perennial ryegrass, meanvalues of site filling were found to be close to this theoreticalmaximum Due to growth-limiting factors, tiller formation from leaf axillarybuds can be delayed or suppressed entirely. A set of equationsis presented from which site filling and the ratio of new leavesto new tillers can be calculated for all situations of axillarybud activity. It is stressed that leaf appearance rate can onlybe determined by marking and counting leaves on single tillersat consecutive dates Loltum perenne (L.), perennial ryegrass, tillering, axillary bud, leaf appearance rate, prophyll, site filling     

Environmental Control of Flowering in Barley (Hordeum vulgare L.). I. Photoperiod Limits to Long-day Responses, Photoperiod-insensitive Phases and Effects of Low-temperature and Short-day Vernalization

Barley plants were grown at a mean diurnal temperature of 15°C and reciprocally transferred between different photoperiods(from 16 h d^–1 to 8, 10 or 13 h d^–1 or vice versaat 4, 8, 16 or 32 d after germination). Ten contrasting genotypeswere examined, including seven spring-sown types-Mona, BGS T16-2,Athenais, Emir, Funza, USDA-016525 and S-37, and three autumn-sowntypes-Gerbel B, Arabi Abiad and Ager. In the latter two alltreatments were repeated on plants grown from seeds which hadbeen vernalized at 2 °C for 42 d. The results suggest that, between the critical photoperiod (belowwhich there is a delay in flowering) and the ceiling photoperiod(below which there is no further delay), there is a linear relationbetween photoperiod and the reciprocal of the time taken toflower (awn emergence). In all genotypes the ceiling photoperiodwas     

Inflorescence Initiation and Leaf Size in Some Gramineae

The morphology of successive leaves on the flowering shoot wasstudied in species of Glyceria, Lolium, and Triticum. The bladesof successive leaves were progressively longer, eventually reachinga maximum, after which the blades of the last few leaves producedbefore heading were shorter. When the longest leaf blade waselongating, dissection of the shoot apices showed that inflorescenceinitiation was taking place. Epidermal cell measurements inTriticum indicate that differences in blade length are due todifferences in the amount of cell extension. It appears that a correlated change occurs in blade morphologyassociated with the onset of the reproductive state of the shootapex, brought about via changes in the amount of cell extension. A study of the effect of different amounts of low-temperatureand different day-lengths on the relation between inflorescenceinitiation and the production of the longest leaf blade showedthat, under some conditions, this relation can be disturbed.     

The Control of Leaf and Ear Size in Barley

The relative growth rates, growth rates, and final size of tillersand main-shoot leaves, internodes, and ear of a freely tilleringspring barley genotype were measured and compared with thoseof a non-tillering single-gene mutant. Leaf and internode growthand final size were greater in the non-tillering mutant. Thedifferences, it is proposed, arise because of changes in internalcompetition for assimilates brought about by the absence oftillers. There was little difference in ear growth or size,possibly because of abnormalities of ear development, whichresulted in fewer spikelets in the non-tillering genotype.