Molecular genetics of growth and development in Populus (Salicaceae). v. mapping quantitative trait loci affecting leaf variation

We examined the genetic variation of leaf morphology and development in the 2-yr-old replicated plantation of an interspecific hybrid pedigree of Populus trichocarpa T. & G. and P. deltoides Marsh. via both molecular and quantitative genetic methods. Leaf traits chosen were those that show pronounced differences between the original parents, including leaf size, shape, orientation, color, structure, petiole size, and petiole cross section. Leaves were sampled from the current terminal, proleptic, and sylleptic branches. In the F2 generation, leaf traits were all significantly different among genotypes, but with significant effects due to genotype X crown-position interaction. Variation in leaf pigmentation, petiole length. And petiole length proportion appeared to be under the control of few quantitative trait loci (QTLs). More QTLs were associated with single leaf area, leaf shape, lamina angle, abaxial color, and petiole flatness, and in these traits the number of QTLs varied among crown positions. In general, the estimates of QTL numbers from Wright's biometric method were close to those derived from molecular markers. For those traits with few underlying QTLs, a single marker interval could explain from 30 to 60% of the observed phenotypic variance. For multigenic traits, certain markers contributed more substantially to the observed variation than others. Genetic cluster analysis showed developmentally related traits to be more strongly associated with each other than with unrelated traits. This finding was also supported by the QTL mapping. For example, the same chromosomal segment of linkage group L seemed to account for 20% of the phenotypic variation of all dimension-related traits, leaf size, petiole length. and midrib angle. In both traits. the P. deltoides alleles had positive effects and were dominant to the P. trichocarpa alleles. Similar relationships were also found for lamina angle. abaxial greenness, and petiole.     

ORGANIZATION AND ONTOGENY OF THE VASCULAR SYSTEM IN THE PETIOLE OF EASTERN COTTONWOOD

The topologic arrangement of petiolar bundles varies within the length of the cottonwood petiole. Each petiolar bundle is formed by the subdivision and aggregation of acropetally differentiating subsidiary bundles in a predictable pattern. The subsidiary bundles provide vascular continuity between the stem and specific portions of the leaf lamina. Spot-labeling of individual veins with ¹⁴CO₂, freeze substitution, and microautoradiography were used to establish the relation between the secondary veins of the lamina and the vasculature of the petiole. Within the petiole vasculature each subsidiary bundle was continuous with a specific portion of the lamina and seemed to have a separate function. Subsidiary bundles continuous with the central leaf trace were closely related functionally to the tip region of the lamina, while the subsidiary bundles continuous with the lateral leaf traces were functionally related to the middle and basal portions of the lamina.     

林缘和荒草坡不同草本层盖度小生境中积雪草的等级可塑性

为研究克隆植物在不同生境中对小生境的利用能力和对不同尺度异质性生境的等级可塑性及其格局差异,比较了林缘和荒草坡不同草本层盖度小生境中积雪草(Centella asiatica)分株种群、克隆片段和叶水平的形态和生理特征。在分株种群水平,林缘和荒草坡积雪草被测的分株种群特征差异均不显著,不同草本层盖度小生境间积雪草根冠比差异显著,即在盖度小的小生境中其根冠比显著大于盖度大的小生境。在克隆片段水平,林缘和荒草坡生境显著影响积雪草的根干重、花果干重、一级匍匐茎节间长和总匍匐茎节间长,而不同草本层盖度小生境则显著影响花果干重、根冠比、一级匍匐茎数、二级匍匐茎节间长和总匍匐茎节间长。荒草坡积雪草拥有更大的根干重和花果干重,但一级匍匐茎节间长和总匍匐茎节间长较短。草本层盖度小的小生境中积雪草的根冠比和一级匍匐茎数显著高于盖度大的小生境,而花果干重、二级匍匐茎节间长和总匍匐茎节间长则相反。在叶水平,荒草坡和草本层盖度大的小生境中,积雪草的叶片长、叶片宽和叶面积都较大,而草本层盖度大的小生境中积雪草叶柄长和比叶柄长都显著大于盖度小的小生境;林缘和荒草坡生境与不同草本层盖度小生境对4个叶绿素指标影响均不显著,林缘和荒草坡生境与不同草本层盖度小生境的交互作用对所有测量特征均无显著影响。上述结果表明,积雪草对不同生境和有草本层盖度差异的小生境的反应存在等级可塑性,但是二者的等级差异格局不同;等级反应格局的差异反映出积雪草对不同生境条件有不同的生态适应对策,且对生境小尺度异质性的可塑性强于对大尺度异质性的可塑性。     

CYCLIC HETEROPHYLLY IN SYNGONIUM (ARACEAE)

Species in the genus Syngonium germinate on the ground and mature on the trunks of the trees. These vines consist of relatively unbranched shoots which grow through the forest for considerable distances both horizontally and vertically. In the three species of this study, the shape of the segment, measured as internode diam/length, falls into two distinct classes, leafy and elongate, and varies cyclicly along single shoots. Two cycles are recognized. One cycle occurs in small diam terrestrial shoots in which cycling seems to be controlled by endogenous factors, and occurs with a period of a few tens of segments. The other cycle occurs in shoots of larger diam which climb and descend from trees. In this cycle, alternation between the two forms is controlled by gain or loss of contact with trees, and shoots may remain within a phase of the cycle for hundreds of segments, as long as the substrate does not change.     

Variation in growth of overwintered stolons of contrasting white clover populations in response to temperature, photoperiod and spring environment

The response of overwintered stolons of nine contrasting white clover populations to temperature, photoperiod and natural conditions was studied in six environments during the spring. Rate of leaf appearance, leaflet length, petiole length, stolon internode length, dry matter distribution within the plant and total dry weight were measured on 15 plants of each population/environment combination. Most characters, except leaf size and proportion of dry matter allocated to leaf, responded positively to temperature in the range 10 – 20°C. A positive effect of photoperiod extension was also found for all characters except rate of leaf appearance, internode length and distribution of dry matter to leaf. Population differences in response to environment were found which were related to both leaf size classification and origin. Stolon dry weight was positively correlated with leaf length, petiole length and stolon internode length in most environments. The relationships between the eight characters were often complex and canonical variate analysis provided a convenient way to discriminate between the populations based on all eight characters.     

THE OCCURRENCE OF INTERCALARY AND UNINTERRUPTED MERISTEMS IN THE INTERNODES OF TROPICAL MONOCOTYLEDONS

The distribution of percent of dividing nuclei, parenchyma cell length, total cell number per internode, and total internode length were determined for successive internodes in the apex and growing vegetative internodes of 23 tropical species in 17 families of monocotyledons. Basal intercalary meristems (IM) were found in representatives of Commelinaceae, Cyperaceae, Flagellariaceae, Poaceae, Restionaceae, and Marantaceae. Uninterrupted meristems (UM) which are confined progressively to the upper region of the internode and are not isolated meristematic regions were found in the Costaceae, Dioscoreaceae, Philesiaceae, Smilacaceae, Agavaceae, Araceae, Arecaceae, Liliaceae, Pandanaceae, and Zingiberaceae. Both IM and UM were found in different species of Orchidaceae. The only morphological trait correlated with meristem type was presence of sheathing leaf bases in all species with IM. Both IM and UM are interpreted as extensions of the primary elongating meristem; the IM is disjunct, and the UM is continuous with it. The phytomer growth unit and the presence of internodal IM's cannot be applied generally to the monocotyledons.     

Construction of genetic linkage map of the medicinal and ornamental plant <Emphasis Type="Italic">Catharanthus roseus</Emphasis>

An integrated genetic linkage map of the medicinal and ornamental plant Catharanthus roseus, based on different types of molecular and morphological markers was constructed, using a F₂ population of 144 plants. The map defines 14 linkage groups (LGs) and consists of 131 marker loci, including 125 molecular DNA markers (76 RAPD, 3 RAPD combinations; 7 ISSR; 2 EST-SSR from Medicago truncatula and 37 other PCR based DNA markers), selected from a total of 472 primers or primer pairs, and six morphological markers (stem pigmentation, leaf lamina pigmentation and shape, leaf petiole and pod size, and petal colour). The total map length is 1131.9 cM (centiMorgans), giving an average map length and distance between two markers equal to 80.9 cM and 8.6 cM, respectively. The morphological markers/genes were found linked with nearest molecular or morphological markers at distances varying from 0.7 to 11.4 cM. Linkage was observed between the morphological markers concerned with lamina shape and petiole size of leaf on LG1 and leaf, stem and petiole pigmentation and pod size on LG8. This is the first genetic linkage map of C. roseus.     

Functional design space of single-veined leaves: role of tissue hydraulic properties in constraining leaf size and shape

BACKGROUND AND AIMS: Morphological diversity of leaves is usually quantified with geometrical characters, while in many cases a simple set of biophysical parameters are involved in constraining size and shape. One of the main physiological functions of the leaf is transpiration and thus one can expect that leaf hydraulic parameters can be used to predict potential morphologies, although with the caveat that morphology in turn influences physiological parameters including light interception and boundary layer thickness and thereby heat transfer and net photosynthesis. METHODS: An iterative model was used to determine the relative sizes and shapes that are functionally possible for single-veined leaves as defined by their ability to supply the entire leaf lamina with sufficient water to prevent stomatal closure. The model variables include the hydraulic resistances associated with vein axial and radial transport, as well as with water movement through the mesophyll and the leaf surface. KEY RESULTS: The four parameters included in the model are sufficient to define a hydraulic functional design space that includes all single-veined leaf shapes found in nature, including scale-, awl- and needle-like morphologies. This exercise demonstrates that hydraulic parameters have dissimilar effects: surface resistance primarily affects leaf size, while radial and mesophyll resistances primarily affect leaf shape. CONCLUSIONS: These distinctions between hydraulic parameters, as well as the differential accessibility of different morphologies, might relate to the convergent evolutionary patterns seen in a variety of fossil lineages concerning overall morphology and anatomical detail that frequently have evolved in linear and simple multi-veined leaves.     

不同荷花品种出泥部分与根状茎生长性状和部分生理指标的比较及相关性分析

在盆栽条件下,对不同荷花(Nelumbo nucifera Gaertn.)品种〔藕莲品种'大紫红'('Dazihong')、花莲品种'中国红·上海'('Zhongguohongshanghai')和花藕兼用莲品种'逸仙莲'('Yixianlian')〕出泥部分(叶和花)和泥下部分(根状茎)的生长性状和部分生理指标进行了比较研究,并对3个荷花品种出泥部分与根状茎间的生长性状和部分生理指标进行了相关性分析.结果表明:'大紫红'在整个生长发育期不开花;'中国红·上海'开花时间相对较早(6月上旬),花重瓣且色泽鲜艳;'逸仙莲'开花时间相对较晚(7月上中旬),花单瓣呈白色.3个荷花品种中,'大紫红'立叶数最少,立叶面积最大,根状茎膨大明显,根状茎中可溶性糖和可溶性蛋白质含量最高;'中国红·上海'立叶数最多,立叶中叶绿素含量最高,根状茎节间长度最长,但根状茎膨大不明显;'逸仙莲'立叶数居中,叶柄最长,根状茎膨大明显,根状茎中可溶性蛋白质、可溶性糖、淀粉和维生素C含量介于其他2个荷花品种之间.相关性分析结果表明:荷花立叶面积与根状茎节间长度呈显著负相关,而与根状茎中可溶性糖、可溶性蛋白质及维生素C含量呈极显著或显著正相关;叶柄长度与根状茎直径和根状茎节间质量呈显著正相关;立叶中叶绿素含量与根状茎节间长度和根状茎中淀粉含量呈显著正相关,而与根状茎中可溶性糖含量呈显著负相关.研究结果显示:根据供试3个荷花品种的立叶面积、叶柄长度、立叶中叶绿素含量可间接判断根状茎的生长发育及营养品质状况,并可进一步应用于快速选育花藕兼用莲品种.     

外界支持物直径对苦瓜生长和觅食行为的影响

采用实验生态学的方法研究了外界支持直径变化对苦瓜植株生长和觅食行为的影响。结果表明,支持物直径变化对主茎节间、叶片、叶柄和卷须的发育进程无明显影响。苦瓜植株的攀援生长显著受到支持物直径大小的影响,当支持物直径小于8mm时,植株能较好地自主攀援生长。生长于直径支持物上的植株比生长于小直径支持物上的植株表现出较高的分枝率、较短的主茎、较长的分枝、较大的比茎长和比叶柄长。相关分析表明,节间长度和叶柄长度同植株自主攀援程度成显著正相关关系,而单叶面积、分枝数量和比叶柄长同植株自主攀援程度成显著负相关关系。作为攀援植物的一种重要资源,外界支持物的特征和存在状况将对植物的生长和行为产生重要影响。     

The functional morphology of light capture and carbon gain in the Redwood forest understorey plant Adenocaulon bicolor Hook

1. A three-dimensional geometric simulation model of crown architecture was utilized to investigate the efficiency of light capture and its relationship to whole-plant CO₂ assimilation of Adenocaulon bicolor .
2. Positioning of the leaves by the combined effects of ontogenetic variations in petiole length and angle and leaf size, and the leaf divergence angles were shown to be effective in minimizing self shading. The efficiency of light absorption varied from 0·64 to 0·70 among individual plants that were sampled.
3. Plant to plant variation in simulated daily carbon gain was strongly influenced by variations in the direct and diffuse PFD received by the individual plants. When simulations were run for all plants under a single common light environment, the carbon gain was strongly dependent on the efficiencies of light absorption of the different plants.
4. Simulations in which petiole length was varied showed a non-linear dependence of light absorption efficiency on petiole length. When both petiole length and leaf size were varied in a way that maintained a constant biomass then an optimal petiole length that corresponded to the observed petiole length was apparent. The observed divergence angle between successive leaves also maximized light absorption efficiency as compared to greater or lesser angles, but increases in internode length had no significant effect.
5. The results of this study provide evidence for selection for an 'optimal design' of crown architecture in Adenocaulon bicolor that maximizes light capture.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

匍匐茎草本金戴戴对盐分梯度的表型可塑性

 研究了匍匐茎型克隆草本金戴戴(Halerpestes ruthenica) 4种基株（基因型）对不同盐分处理（0,85.5, 171.0, 256.5和342.0 mM NaCl）的表型可塑性。随着盐分浓度的增加,实验植物与生长相关的性状指标 (如植株干重、总叶面积、分株数和总匍匐茎长度) 显著减小。植株干重、总叶面积和总匍匐茎长度具有显著的基株间差异。实验植物与形态相关的性状指标 (如平均叶柄长和根冠比) 对盐分梯度具有可塑性并具有显著的基株间差异;而其它形态指标 (如平均节间长、比节间长和比叶柄长)     

Photomorphogenesis of leaves: shade-avoidance and differentiation of sun and shade leaves.

Leaf shape is an important factor in optimal plant growth, because leaves are the main photosynthetic organs. Plants exhibit plasticity in leaf shape and structure, allowing them to optimize photosynthetic efficiency. In Arabidopsis thaliana(L.) Heynh., several types of leaves develop differentially, according to light intensity and quality. When shaded, the expansion of leaf lamina is inhibited, while the petiole elongation is enhanced. This phenomenon is part of the so-called shade-avoidance syndrome. Under low light, A. thaliana develops shade leaves with only one layer of palisade tissue, whereas under high light, it develops sun leaves that have nearly two complete layers of palisade tissue. Although the molecular mechanisms of these photomorphogenic phenomena in leaves are not well understood, recent studies of A. thaliana have provided some insight. For example, some cytochrome P450s may be involved in the specific control of the petiole length during photomorphogenesis. On the other hand, switching between sun and shade leaves is regulated by long-distance signaling from mature leaves in Chenopodium album. Here we provide an overview of the mechanisms of photomorphogenesis in leaves based on recent findings.     

COMPARATIVE MORPHOGENESIS OF THE DIMORPHIC LEAVES OF CYAMOPSIS TETRAGONOLOBA

Cyamopsis tetragonoloba, guar, produces two leaf forms: a simple leaf and a trifoliate leaf. The steps in the development of each of these forms have been investigated in an attempt to determine the precise point at which the leaf primordium becomes destined to produce one or the other characteristic leaf shapes. Up to 140 μ in length the leaf primordia are morphologically indistinguishable. If a simple leaf is to be formed the marginal meristem remains continuous and initiates only lamina. If a trifoliate leaf is to be formed the continuity of the marginal meristem is interrupted by a group of “pocketal” cells dividing it into an upper lamina meristem and a basal leaflet buttress.     

优良单株家系辣木叶的表型性状分析

为挖掘辣木(Moranga oleifera)优良种质资源,对30个优良单株家系的叶片表型性状进行研究。结果表明,除叶形外,辣木不同家系间的叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽均存在不同程度的差异。复叶数与复叶柄长度和直径、复叶间距、叶长、叶宽呈极显著正相关;主成分分析表明,叶长、叶宽、复叶柄长度和直径、复叶间距、叶柄和叶片颜色是区分辣木不同家系最主要的叶片性状指标。聚类分析结果表明,30个辣木家系可分为3大类,叶片表型性状存在显著差异的家系的遗传距离较远。因此,叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽将为直观区分辣木家系提供参考。     

AN INVESTIGATION OF THE PHYLOGENETIC AND ONTOGENETIC VARIABILITY OF SHOOT APICAL MERISTEMS IN THE CACTACEAE

The sizes, shapes and zonations of the shoot apical meristems of 22 species of cacti were examined. This family was chosen because of its great diversity of habits; the more primitive members are nonsucculent. leafy trees and more advanced members are highly specialized “leaf-less” stem-succulents. By combining these measurements with those already in the literature, a sample of almost 70 species was obtained. Apical meristems range in size from only 80 μm in diam in some species to as much as 1.500 μm in diam in others. The shape ranges from being flat to almost hemispherical. Despite the great range in size and shape of the apical meristems, or the range in the morphologies of the leaves and stems which are produced by the meristems. all apices had the usual zonation: tunica, central mother cells, peripheral zone, and pith-rib meristem. The sizes of each of the zones. expressed either as the number of cells per zone or expressed as a percentage of the whole apex. were highly variable. The variation in apical dimensions and zone sizes occurred both phylogenetically and ontogenetically. and this has been interpreted to indicate that the morphogenetic mechanisms which control apical size and zonation are easily modified, both during the development of individual plants and during the evolution of new species.     

Effects of sowing time and plant age on critical nitrogen concentrations in canola (Brassica napus L.)

Critical concentrations of NO₃-N in fresh petiole tissue and total N in the dried lamina were determined for the youngest mature leaf (YML) of field-grown canola. For dry matter yield of canola sown on 4 May, critical NO₃-N concentration in the YML petiole at the rosette stage (RS) was 1.46 mg/g fresh wt. At the flower-buds-visible stage (BV) it was 0.45 mg/g fresh wt. For seed yield the values were 1.72 and 0.53 mg/g fresh wt. Critical total N concentration in the YML lamina for dry matter yield were 69 mg/g dry wt. at RS and 57 at BV. For seed yield they were 71 and 59 mg/g dry wt. Critical NO₃-N concentrations in the YML petiole of canola sown on 30 May were reduced by 50%; critical total-N concentrations in the YML lamina were not reduced to the same extent. Despite the reductions in critical N concentrations in the YML, critical N fertilizer rates for vegetative growth and seed yield were unaffected by sowing date or plant growth stage.     

Petiole mechanics, leaf inclination, morphology, and investment in support in relation to light availability in the canopy of Liriodendron tulipifera

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q_int). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q_int, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q_int. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q_int.