HETEROCHRONY AND HETEROBLASTIC LEAF DEVELOPMENT IN TWO SUBSPECIES OF CUCURBITA ARGYROSPERMA (CUCURBITACEAE)

To date, reports of paedomorphosis at the whole plant or shoot level have been loosely based on whole plant form or on the sequence of leaf shapes produced along the shoot (heteroblasty). However, interpreting the significance of heterochrony in the evolutionary loss or gain of heteroblasty based on mature leaf forms assumes that all leaves with the same shape arose through very similar modes of organogenesis. This study examines this assumption in two subspecies of Cucurbita argyrosperma, one that is wild and heteroblastic and a second that is cultivated and not markedly heteroblastic. All leaves of the cultivar are visually similar to early leaves of the wild subspecies. The cultivar is considered to be the progenitor of the wild subspecies. Scanning electron microscopy and allometry of developing leaves showed that at early nodal positions along the primary shoot, leaf development in both subspecies was similar. At later nodal positions, very young leaves of both subspecies were more similar to each other than to leaves at earlier nodal positions within the same plant at the same stage of development. This early similarity was masked in the mature shapes of later leaves due to subsequent differences in allometric growth. Thus a simple hypothesis of paedomorphosis in which the early leaf form in the progenitor is simply reiterated at later nodal positions in the cultivar is not supported by patterns of leaf development.     

The rates of shoot and root growth in intact plants of pea mutants in leaf morphology

Isogenic lines of pea (Pisum sativum L.) with the genetically determined changes in leaf morphology, afila (af) and tendril-less (tl), were used to study the relationship between shoot and root growth rates. The time-course of shoot and root growth was followed during the pre-floral period in the intact plants grown under similar conditions. The af mutation produced afila leaves without leaflets, whereas in the case of the tl mutations, tendrils were substituted with leaflets, and acacia-like leaves were developed. Due to the changes in leaf morphology caused by these mutations, pea genotypes differed in leaf area: starting from day 7, the leaf area was lower in the af plants and larger in the tl plants as compared to the wild-type plants. Such divergence was amplified in the course of plant development and reached its maximum immediately before the transition to flowering. Plants of isogenic lines did not notably differ in stem surface areas. In spite of significant difference in total leaf area, the wild type and tl plants did not differ in leaf dry weight. Starting from leaf 9, the af plants lagged behind two leaflet-bearing genotypes (wild type and tl) in leaf dry weight, whereas stem dry weight was similar in the wild type and tl forms and slightly lower in the af plants. Root dry weights were practically similar in the wild type and tl plants until flowering. The reduction of leaf area in the af plants drastically reduced root dry weight. In other words, the latter index was related to the total weight and total area of leaves and stems. The correlation analysis demonstrated an extremely low relationship between leaf and stem area and dry weight and those of roots early in plant development (when plants develop five to seven leaves). Later, immediately before flowering (nine to eleven leaves), root weight was positively related to leaf weight and area; however, stem area and root weight did not correlate. Thus, in three genotypes (wild type, af, and tl), at the end of their vegetative growth phase, leaf and root biomass accumulated in proportion, independently of leaf area expansion.     

Developmental events leading to peltate leaf structure in <Emphasis Type="Italic">Tropaeolum majus</Emphasis> (Tropaeolaceae) are associated with expression domain changes of a YABBY gene

Peltate leaf architecture has evolved from conventional bifacial leaves many times in flowering plant evolution. Characteristics of peltate leaves, such as the differentiation of a cross zone and of a radially symmetric, margin-less petiole, have also been observed in mutants of genes responsible for adaxial-abaxial polarity establishment. This suggests that altered regulation of such genes provided a mechanism for the evolution of peltate leaf structure. Here, we show that evolution of leaf peltation in Tropaeolum majus, a species distantly related to Arabidopsis thaliana, was associated with altered expression of Tropaeolum majus FILAMENTOUS FLOWER (TmFIL), a gene conferring abaxial identity. In situ hybridization indicates that adaxial and abaxial domains are established in early leaf primordia as in species with bifacial leaves. Upon initiation of the cross zone by fusion of the blade margins, localized expansion of TmFIL to the upper leaf side could be seen, indicating a local loss of adaxial leaf identity. The observed changes in expression are consistent with a role of TmFIL in radialization of the petiole and circularization of the leaf blade margin by the cross zone. In addition, expression was observed in segment primordia and during expansion of the bifacial blade, suggesting additional roles for TmFIL in leaf development.     

Components of Resistance to Early Blight in Four Potato Cultivars: Effect of Leaf Position

Components of early blight resistance were quantified in leaves of different ages in four potato cultivars. The components of resistance: incubation period (IP), lesion number (LN), early blight severity, lesion expansion rate (LER), latent period (LP) and spore production by lesion area (SPLA), were evaluated separately in the lower, middle and upper leaves of four potato cultivars. Plants of cultivar Aracy (resistant), Delta (moderately resistant), Desirée (susceptible) and Bintje (susceptible) were inoculated with an Alternaria solani isolate at the beginning of the flowering stage. Disease severity varied in different plant parts. In all cultivars, regardless of resistance, the smallest values of LN, and severity were recorded on the upper leaves, suggesting that young tissues are less susceptible. In cultivar Aracy, the IP was long, with small values of LN and LER and consequently, low values of early blight severity in all leaf positions were recorded. Although IP was long in cultivar Aracy, no differences between the moderately resistant cultivar Delta and the susceptible cultivars Bintje and Desirée could be detected for this component. The IP was only influenced by leaf position in cultivar Aracy. Clear differences in resistance levels among cultivars could be detected regarding LN, severity and LER. However, neither LP nor SPLA were associated with resistance level of cultivars or with leaf position. Analyses according to plant part suggest that evaluations on leaves of the middle third part are most suitable for screening for early blight resistance in potato.     

Canopy development of a determinate and an indeterminate cultivar of Vicia faba L. under contrasting plant distributions and densities

Field experiments were conducted in 1987 and 1988 to quantify differences in canopy formation between an indeterminate and a determinate genotype of Vicia faba L., grown at two plant densities and three spatial distributions. The number of stems per unit area produced by determinate plants was related to the growth rate before flowering. Leaf production per stem per unit of thermal time was similar in both plant types, but twice as many leaves per stem were produced by the indeterminate cultivar. The indeterminate cultivar produced fewer and smaller leaves in the warmer and drier weather of 1988 than in 1987. The determinate genotype produced similar sizes and numbers of leaves in both years, but fewer tillers developed in 1988 than in 1987. Accordingly, leaf mass per unit ground area was greater in 1987 than in 1988 in both genotypes. Except during early flowering, relationships between leaf mass and leaf area were constant, with higher specific leaf areas in the determinate than the indeterminate genotype. Shoot dry matter partitioning into leaves was identical in both years for indeterminate plants, but differed in determinate ones.
It is concluded that canopy development is regulated through individual leaf weight and leaf number per stem in non-tillering indeterminate, and by stem numbers per unit area in tillering determinate plants.     

Evolution of salamander life cycles: a major-effect quantitative trait locus contributes to discrete and continuous variation for metamorphic timing

The evolution of alternate modes of development may occur through genetic changes in metamorphic timing. This hypothesis was examined by crossing salamanders that express alternate developmental modes: metamorphosis vs. paedomorphosis. Three strains were used in the crossing design: Ambystoma tigrinum tigrinum (Att; metamorph), wild-caught A. mexicanum (Am; paedomorph), and laboratory Am (paedomorph). Att/Am hybrids were created for each Am strain and then backcrossed to their respective Am line. Previous studies have shown that a dominant allele from Att (met(Att)) and a recessive allele from lab Am (met(lab)) results in metamorphosis in Att/Am hybrids, and met(Att)/met(lab) and met(lab)/met(lab) backcross genotypes are strongly associated with metamorphosis and paedomorphosis, respectively. We typed a molecular marker (contig325) linked to met and found that met(Att)/met(lab) and met(Att)/met(wild) were associated with metamorphosis in 99% of the cases examined. However, the frequency of paedomorphosis was 4.5 times higher for met(lab)/met(lab) than for met(wild)/met(wild). We also found that met(Att)/met(wild) and met(wild)/met(wild) genotypes discriminated distributions of early and late metamorphosing individuals. Two forms of phenotypic variation are contributed by met: continuous variation of metamorphic age and expression of discrete, alternate morphs. We suggest that the evolution of paedomorphosis is associated with genetic changes that delay metamorphic timing in biphasic life cycles.     

Growing larger with domestication: a matter of physiology,morphology or allocation?

• Domestication might affect plant size. We investigated whether herbaceous crops are larger than their wild progenitors, and the traits that influence size variation.
• We grew six crop plants and their wild progenitors under common garden conditions. We measured the aboveground biomass gain by individual plants during the vegetative stage. We then tested whether photosynthesis rate, biomass allocation to leaves, leaf size and specific leaf area (SLA) accounted for variations in whole‐plant photosynthesis, and ultimately in aboveground biomass.
• Despite variations among crops, domestication generally increased the aboveground biomass (average effect +1.38, Cohen's d effect size). Domesticated plants invested less in leaves and more in stems than their wild progenitors. Photosynthesis rates remained similar after domestication. Variations in whole‐plant C gains could not be explained by changes in leaf photosynthesis. Leaves were larger after domestication, which provided the main contribution to increases in leaf area per plant and plant‐level C gain, and ultimately to larger aboveground biomass.
• In general, cultivated plants have become larger since domestication. In our six crops, this occurred despite lower investment in leaves, comparable leaf‐level photosynthesis and similar biomass costs of leaf area (i.e. SLA) than their wild progenitors. Increased leaf size was the main driver of increases in aboveground size. Thus, we suggest that large seeds, which are also typical of crops, might produce individuals with larger organs (i.e. leaves) via cascading effects throughout ontogeny. Larger leaves would then scale into larger whole plants, which might partly explain the increases in size that accompanied domestication.

     

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.     

Cranial differentiation and evolution in Thrichomys apereoides (Rodentia: Echinyidae)

Thrichomys apereoides is an echimyid rodent which ranges in distribution from north-eastern and central Brazil into Paraguay, and currently five subspecies are recognized. Recent morphometric analyses of population samples formally assignable to T. a. laurentius and T. a. inermis , which occur in north-eastern Brazil, have shown that a major group of populations including both subspecies differ in cranial shape from a single population allocated to T. a. laurentius . In this study we employed mathematical models of evolutionary quantitative genetics to assess the role that random drift and selection may have played in the evolution of cranial shape differences in T. apereoides . The hypothesis of evolution due to drift was rejected and the selective forces necessary to account for shape differences were estimated. Minimum selective mortalities of the order of 10^-3 of per generation were sufficient to explain the observed morphologic differentiation.     

Distinct genetic architectures for male and female inflorescence traits of maize

We compared the genetic architecture of thirteen maize morphological traits in a large population of recombinant inbred lines. Four traits from the male inflorescence (tassel) and three traits from the female inflorescence (ear) were measured and studied using linkage and genome-wide association analyses and compared to three flowering and three leaf traits previously studied in the same population. Inflorescence loci have larger effects than flowering and leaf loci, and ear effects are larger than tassel effects. Ear trait models also have lower predictive ability than tassel, flowering, or leaf trait models. Pleiotropic loci were identified that control elongation of ear and tassel, consistent with their common developmental origin. For these pleiotropic loci, the ear effects are larger than tassel effects even though the same causal polymorphisms are likely involved. This implies that the observed differences in genetic architecture are not due to distinct features of the underlying polymorphisms. Our results support the hypothesis that genetic architecture is a function of trait stability over evolutionary time, since the traits that changed most during the relatively recent domestication of maize have the largest effects.     

Structural analysis of the dissected ensiform leaves and shoot morphology of Marathrum foeniculaceum (Podostemaceae)

Foliage leaves of Marathrum foeniculaceum Humb. & Bonpl. (Podostemaceae–Podostemoideae) resemble pinnate compound leaves at first sight. But the similarity is deceptive, since studies of the leaf structure reveal the ensiform (sword-like) shape of the blade. The ribbon-shaped central rachis-like portion represents the parallel-veined ensiform blade that extends in the median plane direction. Pinnae, typically developing from the leaf margin of the blade in transverse position relative to the mother shoot axis, do not occur here, but instead, bi-pinnate-like appendages arise alternately at the adaxial and abaxial edge of the ensiform blade. The pinnate-like structures are accessory structures. These project only from one side (“front side”) of the ensiform blade. The appendages produce repeatedly forked threads that end in pubescent filaments. The trichomes are of the Zeylanidium olivaceum type. The occurrence of similar structures in species of Marathrum and the related genera Apinagia and Mourera are discussed.     

Evolutionary and developmental studies of unifacial leaves in monocots: Juncus as a model system

An important objective in evolutionary developmental biology is to understand the molecular genetic mechanisms that have given rise to morphological diversity. Leaves in angiosperms generally develop as a flattened structure with clear adaxial–abaxial polarity. In monocots, however, a unifacial leaf has evolved in a number of divergent species, in which leaf blades consist of only the abaxial identity. The mechanism of unifacial leaf development has long been a matter of debate for comparative morphologists. However, the underlying molecular genetic mechanism remains unknown. Unifacial leaves would be useful materials for developmental studies of leaf-polarity specification. Moreover, these leaves offer unique opportunities to investigate important phenomena in evolutionary biology, such as repeated evolution or convergent evolution of similar morphological traits. Here we describe the potential of unifacial leaves for evolutionary developmental studies and present our recent approaches to understanding the mechanisms of unifacial leaf development and evolution using Juncus as a model system.     

PHANTASTICA regulates development of the adaxial mesophyll in Nicotiana leaves

Initiation and growth of leaf blades is oriented by an adaxial/abaxial axis aligned with the original axis of polarity in the leaf primordium. To investigate mechanisms regulating this process, we cloned the Nicotiana tabacum ortholog of PHANTASTICA (NTPHAN) and generated a series of antisense transgenics in N. sylvestris. We show that NSPHAN is expressed throughout emerging blade primordia in the wild type and becomes localized to the middle mesophyll in the expanding lamina. Antisense NSPHAN leaves show ectopic expression of NTH20, a class I KNOX gene. Juvenile transgenic leaves have normal adaxial/abaxial polarity and generate leaf blades in the normal position, but the adaxial mesophyll shows disorganized patterns of cell division, delayed maturation of palisade, and ectopic reinitiation of blade primordia along the midrib. Reversal of the phenotype with exogenous gibberellic acid suggests that NSPHAN, acting via KNOX repression, maintains determinacy in the expanding lamina and sustains the patterns of cell proliferation critical to palisade development.     

DOES SHADE PROLONG JUVENILE DEVELOPMENT? A MORPHOLOGICAL ANALYSIS OF LEAF SHAPE CHANGES IN CUCURBIT A ARGYROSPERMA SUBSP. SORORIA (CUCURBITACEAE)

Several studies have concluded that shade extends the juvenile phase of plant development based on the prolonged production of juvenile-looking leaves along the shoot. Until now, the alternative hypothesis that leaves produced in shade converge in shape with more juvenile leaves through plastic responses of individual leaves has not been investigated. The literature has shown that differences in shape among leaves in a heteroblastic series are manifest very early in development, often at or near inception, whereas divergence in development between sun and shade leaves does not become apparent until considerably later. This study is the first to distinguish between these alternatives by comparing the developmental morphology of young leaves of the heteroblastic plant Cucurbita argyrosperma subsp. sororia. Differences in shapes of mature leaves along the shoot in sun and shade were quantified in terms of leaf area, perimeter, and shape using truss analysis. Developmental morphology from initiation through expansion was examined for representative transition and for later (adult) leaves using scanning electron microscopy and allometry. Determinants of shape established very early in development were the same for leaves at the same position grown in sun and shade. Differences in morphology between sun and shade leaves at the same position did not arise until these leaves reached lamina lengths greater than 1,000 μm. Thus, the less-lobed, more juvenile looking leaf produced at later positions in the shade arose through later developmental responses of individual leaves to shade, rather than through a prolonged phase of juvenile development.     

Leaf toughness changes the effectiveness of larval aggregation in the butterfly Byasa alcinous bradanus (Lepidoptera: Papilionidae)

Two subspecies of the papilionid butterfly Byasa alcinous , B. a. bradanus and B. a. alcinous , have varying degrees of larval aggregation. Early instar larvae of ssp. bradanus always occur in aggregations. To determine the functions of larval aggregation in this subspecies, we examined the effects of leaf toughness on larval performance when caterpillars were reared alone and in aggregations. Newly hatched larvae were reared either individually or in groups of 10 and were fed either tough or tender leaves of Aristolochia debilis . When fed tough leaves, more gregarious larvae survived the first instar. This difference between solitary and aggregated larvae did not occur when caterpillars were fed soft leaves. The effects of aggregation on larval weight and duration were not significant between leaf-toughness treatments. Larval aggregation of B. a. bradanus improves larval survivorship in early instars that use host plants with tough leaves.     

Alternative modes of leaf dissection in monocotyledons

Although a majority of monocotyledons have simple leaves, pinnately or palmately dissected blades are found in four orders, the Alismatales, Pandanales, Dioscoreales and Arecales. Independent evolutionary origins of leaf dissection are indicated by phylogenetic analyses and are reflected in the diversity of mechanisms employed during leaf development. The mechanism of blastozone fractionation through localized enhancement and suppression of growth of the free margin of the leaf primordium occurs in the Araceae and Dioscoreaceae. By contrast, the corrugated, dissected leaves of palms (Arecaceae) develop through a two-step process: first, plications are formed through intercalary growth in a submarginal position and, second, the initially simple leaf blade is dissected through an abscission-like process of leaflet separation. A third mechanism, perforation formation, is employed in Monstera and five related genera of the Araceae. In this mode, discrete patches of cells undergo programmed cell death during lamina development, resulting in formation of open perforations. When perforations are positioned near the leaf margin, mechanical disruption of the thin bridges of marginal tissue results in a deeply pinnatisect blade. Whereas blastozone fractionation defines the early primary morphogenesis phase of leaf development, the other two modes occur later, during the secondary morphogenesis/histogenesis phase. Evolution of these mechanisms presumably has involved recruitment of other developmental programmes into the development of dissected leaves.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 25–44.     

Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot

Effects of the length: width ratio of a leaf blade and petiole length on shoot light capture were studied with computer simulation. Both a larger length: width ratio and longer petiole contributed to larger light capture per unit leaf area due to a reduced aggregation of leaf area around the stem. Other conditions being equal, shoots with narrow leaves and no petioles and those with wide leaves with petioles showed similar light capture as long as the mean distance of the leaf blade from the stem was the same. In shoots with a short internode and/or distichous phyllotaxis, however, narrow leaves contributed more to avoiding mutual shading than wide leaves with petioles. The predominance of light coming from a higher angular altitude also favored narrow leaves. The possible consequences of these results in the adaptive geometry of plant architecture are discussed.