Natural developmental variations in leaf and plant senescence in Arabidopsis thaliana

Leaf senescence is a developmentally regulated process that contributes to nutrient redistribution during reproductive growth and finally leads to tissue death. Manipulating leaf senescence through breeding or genetic engineering may help to improve important agronomic traits, such as crop yield and the storage life of harvested organs. Here, we studied natural variations in the regulation of plant senescence among 16 Arabidopsis thaliana accessions. Chlorophyll content and the proportion of yellow leaves were used as indicator parameters to determine leaf and plant senescence respectively. Our study indicated significant genotype effects on the onset and development of senescence. We selected three late- and five early-senescence accessions for further physiological studies. The relationship between leaf and plant senescence was accession-dependent. There was a significant correlation between plant senescence and the total number of leaves, siliques and plant bolting age. We monitored expression of two senescence marker genes, SAG12 and WRKY53 , to evaluate progression of senescence. Our data revealed that chlorophyll content does not fully reflect leaf age, because even fully green leaves had already commenced senescence at the molecular level. Integrating senescence parameters, such as the proportion of senescent leaves, at the whole plant level provided a better indication of the molecular status of the plant than single leaf senescence parameters.     

Leaf lifespan is positively correlated with periods of leaf production and reproduction in 49 herb and shrub species

Leaf life span and plant phenology are central elements in strategies for plant carbon gain and nutrient conservation. Although few studies have found that leaf life span correlate with the patterns of leaf dynamics and reproductive output, but there have not been sufficient conclusive tests for relationships between leaf life span and plant phenological traits, the forms and strengths of such relationships are poorly understood. This study was conducted with 49 herb and shrub species collected from the eastern portion of the Tibetan Plateau and grown together in a common garden setting. We investigated leaf life span, the periods of leaf production and death, the time lag between leaf production and death, and the period of plant reproduction (i.e., flowering and fruiting). Interspecific relationships of leaf life span with leaf dynamics and reproduction period were determined. Leaf production period was far longer than leaf death period and largely reflected the interspecific variation of leaf life span. Moreover, leaf life span was positively correlated with the length of reproduction (i.e., flowering and fruiting) period. These relationships were generally consistent across different subgroups of species (herbs vs. shrubs) and indicate potentially widely applicable relationships between LLS and aboveground phenology. We concluded that leaf life span is associated not simply with the dynamics of the leaf itself but with reproduction period. The results demonstrate a plant trade‐off in resource allocation between production and reproduction and a coordinated arrangement of leaves, flowers, and fruits in their time investment. Our results provide insight into the relationship between leaf life span and plant phenology.     

The role of sugars in integrating environmental signals during the regulation of leaf senescence

Although leaf senescence results in a loss of photosynthetic carbon fixation, the senescence-dependent release of nutrients, especially of nitrogen, is important for the growth of young leaves and for reproduction. Environmental regulation of senescence is therefore a vital factor in the carbon and nitrogen economy of plants. Leaf senescence is a highly plastic trait that is affected by a range of different environmental factors including light, nutrient supply, CO2 concentration, and abiotic and biotic stress. In this review, the focus is on the impact of environmental conditions on sugar accumulation and sugar signalling during senescence. By signalling a high availability of carbon relative to nitrogen in the old leaves, sugar accumulation can trigger leaf senescence. Sugar-induced senescence is therefore particularly important under low nitrogen availability and may also play a role in light signalling. Whether or not sugars are involved in regulating the senescence response of plants to elevated CO2 remains unresolved. Senescence can be delayed or accelerated in elevated CO2 and no clear relationship between sugar accumulation and senescence has been found. Plasticity in the response to environmental factors, such as daylength and sugar accumulation, varies between different Arabidopsis accessions. This natural variation can be exploited to analyse the genetic basis of the regulation of senescence and the consequences for growth and fecundity. Different evolutionary strategies, i.e. early senescence combined with a high reproductive effort or late senescence combined with a low reproductive effort, may be an important adaptation of Arabidopsis accessions to their natural habitat.     

Evolutionary potential in the Alpine: trait heritabilities and performance variation of the dwarf willow Salix herbacea from different elevations and microhabitats

Alpine ecosystems are seriously threatened by climate change. One of the key mechanisms by which plants can adapt to changing environmental conditions is through evolutionary change. However, we still know little about the evolutionary potential in wild populations of long‐lived alpine plants. Here, we investigated heritabilities of phenological traits, leaf size, and performance traits in natural populations of the long‐lived alpine dwarf shrub Salix herbacea using relatedness estimates inferred from SSR (Simple Sequence Repeat) markers. Salix herbacea occurs in early‐ and late‐snowmelt microhabitats (ridges and snowbeds), and we assessed how performance consequences of phenological traits and leaf size differ between these microhabitats in order to infer potential for evolutionary responses. Salix herbacea showed low, but significant, heritabilities of leaf size, clonal and sexual reproduction, and moderate heritabilities of phenological traits. In both microhabitats, we found that larger leaves, longer intervals between snowmelt and leaf expansion, and longer GDD (growing‐degree days) until leaf expansion resulted in a stronger increase in the number of stems (clonal reproduction). In snowbeds, clonal reproduction increased with a shorter GDD until flowering, while the opposite was found on ridges. Furthermore, the proportion of flowering stems increased with GDD until flowering in both microhabitats. Our results suggest that the presence of significant heritable variation in morphology and phenology might help S. herbacea to adapt to changing environmental conditions. However, it remains to be seen if the rate of such an evolutionary response can keep pace with the rapid rate of climate change.     

Quantitative trait locus analysis of growth-related traits in a new Arabidopsis recombinant inbred population

Arabidopsis natural variation was used to analyze the genetics of plant growth rate. Screening of 22 accessions revealed a large variation for seed weight, plant dry weight and relative growth rate but not for water content. A positive correlation was observed between seed weight and plant area 10 d after planting, suggesting that seed weight affects plant growth during early phases of development. During later stages of plant growth this correlation was not significant, indicating that other factors determine growth rate during this phase. Quantitative trait locus (QTL) analysis, using 114 (F9 generation) recombinant inbred lines derived from the cross between Landsberg erecta (Ler, from Poland) and Shakdara (Sha, from Tadjikistan), revealed QTLs for seed weight, plant area, dry weight, relative growth rate, chlorophyll fluorescence, flowering time, and flowering-related traits. Growth traits (plant area, dry weight, and relative growth rate) colocated at five genomic regions. At the bottom of chromosome 5, colocation was found of QTLs for leaf area, leaf initiation speed, specific leaf area, and chlorophyll fluorescence but not for dry weight, indicating that this locus might be involved in leaf development. No consistent relation between growth traits and flowering time was observed despite some colocations. Some of the QTLs detected for flowering time overlapped with loci detected in other recombinant inbred line populations, but also new loci were identified. This study shows that Arabidopsis can successfully be used to study the genetic basis of complex traits like plant growth rate.     

不同品种(系)小豆根系活力与叶片衰老的关联研究

以高产小豆品种(系)‘2000-75’、‘冀红9218’和低产品种(系)‘红宝1号’、‘湾选1号’为材料,测定小豆开花至成熟期,根系与始花节位叶片生长指标、保护酶活性、可溶性蛋白含量等变化,分析其根系与始花叶协同衰老的相关性,以揭示小豆根系活力与地上部叶片衰老之间的关系。结果表明:(1)各品种(系)开花后,植株根系伤流强度随着花后生育进程的推进呈单峰增长趋势,但根系活力的衰老起始期晚于叶片功能的衰退开始期。(2)从开花至成熟期,小豆根系活力与叶片中超氧化物歧化酶、过氧化氢酶、叶绿素含量、可溶性蛋白、叶片净光合速率都存在正相关关系,与丙二醛含量呈负相关关系。(3)与低产品种相比,高产小豆品种(系)‘2000-75’和‘冀红9218’的根系活力强,叶片功能期持续时间长,叶绿素含量下降速度慢,保护性酶含量高,使花后小豆的功能叶捕获光能的能力增强,从而形成较高光合能力的小豆群体,最终获得高产。     

EFFECTS OF PHOTOPERIOD AND VERNALIZATION ON THE NUMBER OF LEAVES AT FLOWERING IN 32 ARABIDOPSIS THALIANA (BRASSICACEAE) ECOTYPES

The extent of natural variation among wild type Arabidopsis thaliana L. Heynh for response to environmental stimuli that affect flowering is poorly documented. The effects of photoperiod and vernalization on the number of rosette leaves at the time of anthesis was measured for 32 Arabidopsis ecotypes. All ecotypes were vernalized 24 days at 4 C under continuous illumination. Vernalized and nonvernalized plants were transplanted into 8- (short-day) and 20-hour (long-day) photoperiods in controlled environment growth chambers. Two ecotypes failed to flower after 110 days. Mean leaf number was less for all ecotypes under long day compared to short day. Sixteen ecotypes responded to vernalization; eight had reduced leaf number regardless of photoperiod, and eight had reduced leaf number only under short day. Two ecotypes required vernalization to flower in this study. For three ecotypes, vernalization and short day resulted in a reduction in leaf number whereas vernalization and long day resulted in increased leaf number. Documentation of the effects and interactions of photoperiod and vernalization across many ecotypes provides a broader range of described natural variation for genetic and physiologic study.     

Natural variation in the regulation of leaf senescence and relation to N and root traits in wheat

Objectives

To identify parameters that can be used for the analysis of natural variation in leaf senescence of wheat; and to understand the association between the onset and progression of leaf senescence with N uptake and root traits.

Methods

Chlorophyll content and the proportion of yellow leaves were used as senescence indicators and their relation with other morphological and physiological traits were measured in contrasting early senescing (ES) and late senescing (LS) wheat lines.

Results

There were significant genotype effects on the onset and progress of senescence. The ES lines in which leaf senescence commenced early had significantly lower root biomass and N uptake than LS lines. The strong negative association between the extent of leaf senescence with root biomass and N uptake indicated that the poor root growth induced N limitation caused the early senescence of ES lines.

Conclusions

The leaf senescence development in ES lines was precocious and constitutive as the trait expressed even under optimal growth conditions suggesting they could be useful in understanding the genetic regulation of senescence under different abiotic stress situations. Accelerated leaf senescence in wheat could be a mechanism to compensate for limitations in the root system that tend to restrict nutrient uptake.     

GENETICS OF BRASSICA CAMPESTRIS. 1. GENETIC CONSTRAINTS ON EVOLUTION OF LIFE-HISTORY CHARACTERS

Energy allocation arguments suggest a possible tradeoff between timing and magnitude of reproduction: plants that postpone reproduction may accumulate greater resources and consequently produce more offspring. However, early reproduction may be favored when adult mortality is high. Tradeoffs among life-history characters may be a consequence of constraints imposed by genetic and environmental covariation among traits. In this paper we examine the genetic basis of the relationship between timing and magnitude of reproduction in an annual plant, Brassica campestris, by selecting to change flowering date and plant size in each of four directions (early and large, late and large, early and small, or late and small). There is a strong positive relationship between flowering date and flowering height. The response to selection was greatest along the axis of positive genetic covariation. Populations may evolve to become early flowering and small or late flowering and tall, but there is little response for the alternative combinations of characters. In this instance, the constraints imposed by quantitative genetics are in striking accord with predictions that might be made on physiological, energetic, or ecological grounds.     

Environmental conditions and genetic differentiation: what drives the divergence of coexisting Leymus chinensis ecotypes in a large-scale longitudinal gradient?

Aims The two coexisting Leymus chinensis ecotypes exhibit remarkable divergences in adaptive strategies under drought and salinity in semi-humid meadows and semi-arid steppes. In order to detect the major genetic and environmental factors dominating the intraspecific phenotype variations and ecotype formation, the questions regarding the two distinct phenotypic forms (ecotypes) in L. chinensis were addressed: (i) did environments drive the L. chinensis ecotype formation? (ii) was there a molecular basis for the morphological divergence between the two ecotypes? (iii) which driving force dominated the intraspecies divergence, divergent natural selection, genetic drift or stabilizing selection?Methods We applied a series experiments on demographical, morphological and physiological traits of two Leymus chinensis ecotypes with gray green (GG) and yellow green (YG) leaf color in nine wild sites along a longitudinal gradient from 114° to 124°E in northeast China. The environmental data including mean annual precipitation, mean annual temperature, elevation and soil properties were collected. We compared the differences of morphological, physiological and genetic differentiations between the two ecotypes.Important findings The GG type exhibited stronger fitness than YG type from the population densities, morphological traits (e.g. shoot height, leaf area, leaf and seed weights et al.), leaf mass per area (LMA) and physiological traits [relative water content (RWC), proline, soluble sugar contents]. Most of above phenotypes (e.g. total shoot densities, spike length et al.) were significantly correlated with mean annual precipitation, mean annual temperature and soil water content (SWC), rarely a correlated with soil pH and soil nutrient. Transplanted populations showed convergence trend by their leaf chlorophyll contents and osmotic adjustments (proline and soluble sugar contents) in the greenhouse, but still exhibited their divergences between two ecotypes in the outdoor transplantation, suggesting that whether L. chinensis ecotype differentiated could be largely affected by the environmental conditions. Furthermore, by the comparison result of quantitative genetic variation (Q ST) values from phenotypes with theoretical neutral genetic differentiation (F ST), differentiation in phenotypic traits greatly surpassed neutral predictions, implying that directional natural selection played a crucial role in L. chinensis ecotype differentiation. In addition, microsatellite analysis from Neighbor-joining tree and Bayesian assignment generated into two groups according to ecotypes, indicating molecular genetic differentiation also propelled the two ecotypes divergence. We conclude that L. chinensis population variations were driven by combing divergent natural selection (precipitation, temperature and SWCs) along the large-scale gradient and significantly intrinsic genetic differentiation.     

Patterns of ehtylene production in senescing leaves

Changes in the patterns of ethylene production, chlorophyll content, and respiration were studied in relation to the senescence of intact leaves and leaf discs. The primary leaves of pinto bean, which abscise readily during natural senescence, and tobacco and sugar beet leaves, which do not abscise, were used. A decrease in the rate of ethylene production and respiration, during the slow phase of chlorophyll degradation, was observed in leaf-blade discs cut from mature leaves and aged in the dark. During rapid chlorophyll loss both ethylene production and respiration increased and then decreased. These climacteric-like patterns were shown by leaf discs of all three species. Discs taken from leaves that had been senescing on the plant also showed a climacteric-like rise in ethylene production but not in respiration, which decreased continuously with leaf age. Climacteric-like patterns in the rise of ethylene and respiration for leaf discs were also shown by the petioles of both bean and tobacco leaves. This indicates that the rise of ethylene and respiration is characteristic of the general process of senescence in leaves and is not restricted to the abscission process. In contrast to the ethylene-forming systems in climacteric fruits and many flowers, the one in leaves declines sharply in the early stages of senescence. The subsequent rise of ethylene production appears to be associated with the rapid phase of chlorophyll breakdown, and may indicate the final stage of the senescence process during which ethylene could be actively involved in inducing leaf abscission.     

Heritable variation and genetic correlation of quantitative traits within and between ecotypes of Avena barbata

We examined heritable variation for quantitative traits within and between naturally occurring mesic and xeric ecotypes of the slender wild oat (Avena barbata), and in 188 recombinant inbred lines derived from a cross between the ecotypes. We measured a suite of seedling and adult traits in the greenhouse, as well as performance-related traits in field sites native to the two ecotypes. Although the ecotypes were genetically diverged for most traits, few traits showed significant heritable variation within either ecotype. In contrast, considerable heritable variation was released in the recombinant progeny of the cross, and transgressive segregation was apparent in all traits. Heritabilities were substantially greater in the greenhouse than in the field, and this was associated with an increase in environmental variance in the field, rather than a decrease in genetic variance. Strong genetic correlations were evident among the recombinants, such that 22 measured traits could be well represented by only seven underlying factors, which accounted for 80% of the total variation. The primary axis of variation in the greenhouse described a trade-off between vegetative and reproductive allocation, mediated by the date of first flowering, and fitness was strongly correlated with this trade-off. Other factors in the greenhouse described variation in size and in seedling traits. Lack of correlation among these factors represents the release of multivariate trait variation through recombination. In the field, a separate axis of variation in overall performance was found for each year/site combination. Performance was significantly correlated across field environments, but not significantly correlated between greenhouse and field.     

ADAPTATION TO FINE-GRAINED ENVIRONMENTAL VARIATION: AN ANALYSIS OF WITHIN-INDIVIDUAL LEAF VARIATION IN AN ANNUAL PLANT

Recent studies of evolution in heterogeneous environments have concentrated on the role of coarse-grained environmental variation. Here I explore the potential for a modular organism to adapt to fine-grained environmental variation through within-individual variation among modules. I describe the pattern of variation among leaves of single individuals and report results of initial analyses of genetic variation for within-individual variability in leaf traits and of genetic correlations that could influence the rate of further evolution of within-individual variation of these traits. Plants from 24 paternal half-sib families were raised in growth chambers, and five traits were measured for two leaves produced by each plant. Four of the five traits differed significantly between sampling times. Genetic analyses revealed significant additive genetic variation for within-individual variation in several traits. Estimates of family mean correlations between traits expressed at different times suggest few relationships that would be expected to impede response to selection for changes in the pattern of within-individual variation in leaf traits. These results support the possibility that within-individual variation could evolve as an adaptive response to fine-grained environmental variation and suggest a need for further investigation to improve understanding of evolution in heterogeneous environments.     

Correlation of leaf senescence and gene expression/activities of chlorophyll degradation enzymes in harvested Chinese flowering cabbage (Brassica rapa var. parachinensis)

Chinese flowering cabbage is one of the main leafy vegetables produced in China. They have a rapid leaf yellowing due to chlorophyll degradation after harvest that limits their marketing. In the present study, leaf senescence of the cabbages was manipulated by ethylene and 6-benzyl aminopurine (6-BA) treatment to investigate the correlation of leaf senescence and chlorophyll degradation related to gene expression/activities in the darkness. The patterns of several senescence associated markers, including a typical marker, the expression of senescence-associated gene SAG₁₂, demonstrated that ethylene accelerated leaf senescence of the cabbages, while 6-BA retarded this progress. Similar to the trends of BrSAG₁₂ gene expression, strong activation in the expression of three chlorophyll degradation related genes, pheophytinase (BrPPH), pheophorbide a oxygenase (BrPAO) and red chlorophyll catabolite reductase (BrRCCR), was detected in ethylene treated and control leaves during the incubation, while no evident increase was recorded in 6-BA treated leaves. The overall dynamics of Mg-dechelatase activities in all treatments displayed increasing trends during the senescence process, and a delayed increase in the activities was observed for 6-BA treated leaves. However, chlorophyllase activity as well as the expression of BrChlase1 and BrChlase2 decreased with the incubation in all treatments. Taken together, the expression of BrPPH, BrPAO and BrRCCR, and the activity of Mg-dechelatase was closely associated with the chlorophyll degradation during the leaf senescence process in harvested Chinese flowering cabbages under dark conditions.     

Effects of P(SAG12)-IPT gene expression on development and senescence in transgenic lettuce

An ipt gene under control of the senescence-specific SAG12 promoter from Arabidopsis (P(SAG12)-IPT) significantly delayed developmental and postharvest leaf senescence in mature heads of transgenic lettuce (Lactuca sativa L. cv Evola) homozygous for the transgene. Apart from retardation of leaf senescence, mature, 60-d-old plants exhibited normal morphology with no significant differences in head diameter or fresh weight of leaves and roots. Induction of senescence by nitrogen starvation rapidly reduced total nitrogen, nitrate, and growth of transgenic and azygous (control) plants, but chlorophyll was retained in the lower (outer) leaves of transgenic plants. Harvested P(SAG12)-IPT heads also retained chlorophyll in their lower leaves. During later development (bolting and preflowering) of transgenic plants, the decrease in chlorophyll, total protein, and Rubisco content in leaves was abolished, resulting in a uniform distribution of these components throughout the plants. Homozygous P(SAG12)-IPT lettuce plants showed a slight delay in bolting (4-6 d), a severe delay in flowering (4-8 weeks), and premature senescence of their upper leaves. These changes correlated with significantly elevated concentrations of cytokinin and hexoses in the upper leaves of transgenic plants during later stages of development, implicating a relationship between cytokinin and hexose concentrations in senescence.     

A latitudinal cline and response to vernalization in leaf angle and morphology in Arabidopsis thaliana (Brassicaceae)

Adaptation to latitudinal patterns of environmental variation is predicted to result in clinal variation in leaf traits. Therefore, this study tested for geographic differentiation and plastic responses to vernalization in leaf angle and leaf morphology in Arabidopsis thaliana. Twenty-one European ecotypes were grown in a common growth chamber environment. Replicates of each ecotype were exposed to one of four treatments: 0, 10, 20 or 30 d of vernalization. Ecotypes from lower latitudes had more erect leaves, as predicted from functional arguments about selection to maximize photosynthesis. Lower-latitude ecotypes also had more elongated petioles as predicted by a biomechanical constraint hypothesis. In addition, extended vernalization resulted in shorter and more erect leaves. As predicted by functional and adaptive hypotheses, our results show genetically based clinal variation as well as environmentally induced variation in leaf traits.     

Adaptive divergence in flowering time among natural populations of Arabidopsis thaliana: Estimates of selection and QTL mapping

To identify the ecological and genetic mechanisms of local adaptation requires estimating selection on traits, identifying their genetic basis, and evaluating whether divergence in adaptive traits is due to conditional neutrality or genetic trade‐offs. To this end, we conducted field experiments for three years using recombinant inbred lines (RILs) derived from two ecotypes of Arabidopsis thaliana (Italy, Sweden), and at each parental site examined selection on flowering time and mapped quantitative trait loci (QTL). There was strong selection for early flowering in Italy, but weak selection in Sweden. Eleven distinct flowering time QTL were detected, and for each the Italian genotype caused earlier flowering. Twenty‐seven candidate genes were identified, two of which (FLC and VIN3) appear under major flowering time QTL in Italy. Seven of eight QTL in Italy with narrow credible intervals colocalized with previously reported fitness QTL, in comparison to three of four in Sweden. The results demonstrate that the magnitude of selection on flowering time differs strikingly between our study populations, that the genetic basis of flowering time variation is multigenic with some QTL of large effect, and suggest that divergence in flowering time between ecotypes is due mainly to conditional neutrality.     

Latitudinal variation in plant size and relative growth rate in Arabidopsis thaliana

Latitude is an important determinant of local environmental conditions that affect plant growth. Forty ecotypes of Arabidopsis thaliana were selected from a wide range of latitudes (from 16°N to 63°N) to investigate genetic variation in plant size and relative growth rate (RGR) along a latitudinal gradient. Plants were grown in a greenhouse for 31 days, during which period three consecutive harvests were performed. Plants from high latitudes tended to have smaller plant size in terms of seed size, cotyledon width, rosette size, number of rosette leaves, size (leaf area) of the largest leaves, total leaf area, and total dry weight per plant than those from low latitudes. The mean (±SE) RGR across ecotypes was 0.229 (±0.0013) day⁻¹. There was, however, significant ecotypic variation, with RGR being negatively correlated with latitude. The two main components of RGR, leaf area ratio (LAR) and unit leaf rate (ULR), were also correlated with latitude: LAR increased with increasing latitude while ULR decreased with increasing latitude. It was also found that RGR tended to be negatively correlated with LAR, specific leaf area (SLA) and specific root length (SRL) but to be positively correlated with mean area per leaf (MAL) and ULR. The variation in RGR among ecotypes was relatively small compared with that in the other traits. RGR may be a conservative trait, whose variation is constrained by the trade-off between its physiological (i.e. ULR) and morphological (i.e. LAR) components. Received: 2 November 1997 / Accepted: 28 February 1998     

香椿种质生长及叶部表型性状的遗传变异分析

以84个香椿(Toona sinensis(A.Juss.)Roem.)种质为材料,对其2个生长性状和18个叶部性状(包含6个质量性状和12个数量性状)进行测定。结果显示,香椿6个叶部质量性状变异类型丰富,呈现出多态化特点,单一性状的主要表型多为1~2个。苗高、地径及叶部表型等14个数量性状在种质间的差异均达到极显著水平,且除地径外,其他性状的遗传方差分量均大于环境方差分量,表明此类性状主要受遗传控制。参试的14个数量性状的平均表型变异系数为20.35%,平均遗传变异系数为16.36%;综合表型和遗传变异系数,叶柄长度较其他性状变异大,而叶片夹角稳定性最高,各数量性状(除地径外)遗传变异系数与表型变异系数之差小于7%。香椿种质各性状间Shannon-Weaver遗传多样性指数相差不大(1.892~2.069),遗传多样性水平高,具有良好的遗传改良基础。聚类分析可将84个香椿种质分为5类,类群Ⅰ表现为生长旺盛、小比叶重型;类群Ⅱ生长较快、叶片较大;类群Ⅲ种质数量最多,属生长缓慢、大比叶重型;类群Ⅳ特征为大叶片、多叶型;类群Ⅴ为小叶片、稀叶型。研究结果表明参试香椿种质变异丰富,遗传多样性水平高,能为良种选育、遗传改良等研究提供丰富的遗传材料。