Evolution of leaf form correlates with tropical-temperate transitions in Viburnum (Adoxaceae)

Strong latitudinal patterns in leaf form are well documented in floristic comparisons and palaeobotanical studies. However, there is little agreement about their functional significance; in fact, it is still unknown to what degree these patterns were generated by repeated evolutionary adaptation. We analysed leaf form in the woody angiosperm clade Viburnum (Adoxaceae) and document evolutionarily correlated shifts in leafing habit, leaf margin morphology, leaf shape and climate. Multiple independent shifts between tropical and temperate forest habitats have repeatedly been accompanied by a change between evergreen, elliptical leaves with entire margins and deciduous, more rounded leaves with toothed or lobed margins. These consistent shifts in Viburnum support repeated evolutionary adaptation as a major determinant of the global correlation between leaf form and mean annual temperature. Our results provide a new theoretical grounding for the inference of past climates using fossil leaf assemblages.     

Behavioral cues associated with oviposition by Trioza eugeniae

Trioza eugeniae oviposited significantly more eggs on Syzygium paniculatum leaves free of eggs compared to leaves with pre-existing eggs in a choice experiment, suggesting that females modify oviposition based on cues associated with the presence of eggs. In separate experiments, females avoided ovipositing on parts of leaves where eggs were present, but readily oviposited on areas of the same leaf without eggs. Epideictic pheromones are apparently not used by this species because females readily oviposited on areas of a leaf from which eggs had previously been removed. Females laid 57% of all their eggs along the margins of normal leaves, but also oviposited on artificial margins produced by cutting the leaves. The preference for natural margins over artificial margins was eliminated when the entire margin was coated with an antitranspirant compound to mask the damaged edge. These studies show that ovipositing psyllids respond both to the presence of eggs and to the suitability of the substrate.     

Red (anthocyanic) leaf margins do not correspond to increased phenolic content in New Zealand Veronica spp.

Background and Aims

Red or purple coloration of leaf margins is common in angiosperms, and is found in approx. 25 % of New Zealand Veronica species. However, the functional significance of margin coloration is unknown. We hypothesized that anthocyanins in leaf margins correspond with increased phenolic content in leaf margins and/or the leaf entire, signalling low palatability or leaf quality to edge-feeding insects.

Methods

Five species of Veronica with red leaf margins, and six species without, were examined in a common garden. Phenolic content in leaf margins and interior lamina regions of juvenile and fully expanded leaves was quantified using the Folin–Ciocalteu assay. Proportions of leaf margins eaten and average lengths of continuous bites were used as a proxy for palatability.

Key Results

Phenolic content was consistently higher in leaf margins compared with leaf interiors in all species; however, neither leaf margins nor more interior tissues differed significantly in phenolic content with respects to margin colour. Mean phenolic content was inversely correlated with the mean length of continuous bites, suggesting effective deterrence of grazing. However, there was no difference in herbivore consumption of red and green margins, and the plant species with the longest continuous grazing patterns were both red-margined.

Conclusions

Red margin coloration was not an accurate indicator of total phenolic content in leaf margins or interior lamina tissue in New Zealand Veronica. Red coloration was also ineffective in deterring herbivory on the leaf margin, though studies controlling for variations in leaf structure and biochemistry (e.g. intra-specific studies) are needed before more precise conclusions can be drawn. It is also recommended that future studies focus on the relationship between anthocyanin and specific defence compounds (rather than general phenolic pools), and evaluate possible alternative functions of red margins in leaves (e.g. antioxidants, osmotic adjustment).     

Patterns of leaf size and morphology in relation to shoot length in Tsuga canadensis

Summary Pattern of change in leaf character was assessed along the length and around the circumference of Tsuga canadensis (L.) Carr. shoots of different length classes. Leaf size decreased, and number of leaves per unit length of shoot increased, with decrease in shoot length. Acropetally along the shoots, lamina length and width decreased, relative petiole length increased, apices became more pointed, and leaf margins bore more teeth. Around the shoot, from upper to side, and to lower surfaces, leaf size and number of marginal teeth in the proximal halves of leaves increased. These patterns were related both to production of preformed versus neoformed leaves, though their separation was indistinct, and to secondary orientation of leaves by twisting in their petiolar regions into two major bi-lateral ranks. An additional minor rank occurred along the upper surfaces of the shoots where secondary orientation of the leaves was minimal. Surface features of leaves did not differ in any obvious manner. Leaves on sylleptic shoots, which by definition were all neoformed, exhibited similar patterns, but were generally smaller than those on their parent shoots.     

Leaf shape and size track habitat transitions across forest–grassland boundaries in the grass family (Poaceae)

Grass leaf shape is a strong indicator of their habitat with linear leaves predominating in open areas and ovate leaves distinguishing forest‐associated grasses. This pattern among extant species suggests that ancestral shifts between forest and open habitats may have coincided with changes in leaf shape or size. We tested relationships between habitat, climate, photosynthetic pathway, and leaf shape and size in a phylogenetic framework to evaluate drivers of leaf shape and size variation over the evolutionary history of the family. We also estimated the ancestral habitat of Poaceae and tested whether forest margins served as transitional zones for shifts between forests and grasslands. We found that grass leaf shape is converging toward different shape optima in the forest understory, forest margins, and open habitats. Leaf size also varies with habitat. Grasses have smaller leaves in open and drier areas, and in areas with high solar irradiance. Direct transitions between linear and ovate leaves are rare as are direct shifts between forest and open habitats. The most likely ancestral habitat of the family was the forest understory and forest margins along with an intermediate leaf shape served as important transitional habitat and morphology, respectively, for subsequent shifts across forest–grassland biome boundaries.     

Comparative analysis of leaf shape development in Eschscholzia californica and other Papaveraceae-Eschscholzioideae

Dissected leaves in Papaveraceae-Eschscholzioideae have an architecture frequently encountered in the basal eudicot clade Ranunculales that could represent an ancestral condition for eudicots. Developmental morphology of foliage leaves was investigated using scanning electron microscopy and focusing on primordium formation activity (primary morphogenesis) at the leaf margin. Eschscholzia californica, E. lobii, and Hunnemannia fumariaefolia had a polyternate-acropetal mode of leaf dissection. Segment formation continued around the whole leaf blade periphery. Differences in mature leaf architecture was traced to variations in regional blastozone activity and duration. Epidermal cell size measurements in E. californica indicated that the leaf tip tissue starts to differentiate already at the onset of organogenic activity and that tip cells remain larger than epidermal cells at the basal margins during further growth. It is argued that early differentiation of the tip does not set up a general basipetal differentiation gradient, but is a local effect that allows acropetal pinna initiation to occur in subapical blastozones. In Dendromecon, secondarily entire leaves have evolved through the loss of primordium formation activity. Marginal corrugations found in Dendromecon form late in development and are not reminiscent of lateral primordia.     

The influence of leaf size and shape on leaf thermal dynamics: does theory hold up under natural conditions?

Laboratory studies on artificial leaves suggest that leaf thermal dynamics are strongly influenced by the two‐dimensional size and shape of leaves and associated boundary layer thickness. Hot environments are therefore said to favour selection for small, narrow or dissected leaves. Empirical evidence from real leaves under field conditions is scant and traditionally based on point measurements that do not capture spatial variation in heat load. We used thermal imagery under field conditions to measure the leaf thermal time constant (τ) in summer and the leaf‐to‐air temperature difference (?T) and temperature range across laminae (T_range) during winter, autumn and summer for 68 Proteaceae species. We investigated the influence of leaf area and margin complexity relative to effective leaf width (w_e), the latter being a more direct indicator of boundary layer thickness. Normalized difference of margin complexity had no or weak effects on thermal dynamics, but w_e strongly predicted τ and ?T, whereas leaf area influenced T_range. Unlike artificial leaves, however, spatial temperature distribution in large leaves appeared to be governed largely by structural variation. Therefore, we agree that small size, specifically w_e, has adaptive value in hot environments but not with the idea that thermal regulation is the primary evolutionary driver of leaf dissection.     

Larval gregariousness and neonate establishment of the eucalypt-feeding beetle Chrysophtharta agricola (Coleoptera: Chrysomelidae: Paropsini)

The selective advantage offered to individuals living within groups may relate to natural enemy defence, but in leaf feeding insects may also relate to overcoming plant defences, especially with respect to feeding establishment. We conducted a series of experiments focusing on neonate larval survival, examining the effect of group size and leaf age on the survival of a eucalypt-feeding beetle, Chrysophtharta agricola , which formed groups of up to 43 larvae on the foliage of Eucalyptus nitens in the field. In the laboratory, in the absence of natural enemies, we found that initial density, leaf age and damage to the leaf margin significantly affected larval survival. Survival of solitary first-instar larvae on young foliage was around 80% whereas on older foliage it was around 11%. Prior damage to the leaf margin significantly increased survival on older leaves to around 61%. Initial larval density also affected survival, although mortality was always significantly higher on older leaves. On older leaves the larval group size above which mortality increased no further was over two-fold that on young leaves. Observations of group feeding behaviour at each instar showed that the majority of larvae (75.7%) were aligned facing away from the feeding site and that only around 7.5%, or just 1–2 larvae per group, fed at any one time. Feeding larvae chewed the leaf edge by straddling the leaf margin. Measurements of leaf margins showed that older leaves had significantly thicker leaf margins and 'thickness' ratios (leaf margin to leaf lamina proper). In the field, approximately 85% of all larvae occurred on the first two expanded leaf pairs, and larval mortality was highest between eclosion and establishment of the first instar. However, beetles apparently did not adjust clutch size according to leaf age.     

Variation in stomatal characteristics over the lower surface of Commelina communis leaves

Abstract. The silicone rubber impression technique was used to measure stomatal apertures in 9 mm² sampling areas covering the entire lower surface of leaves of Commelina communis L. The data were analysed using a computer program which produced 'iso-aperture' contours illustrating local differences in mean stomatal aperture. Little consistency was seen in the iso-aperture patterns among sampling times, although the stomata were always relatively closed at the leaf tip and base. When stomata in the middle of the lamina were open, those near to the leaf margin tended to be relatively closed. In places, gradients of mean stomatal aperture were as high as 1 μm mm⁻¹. Measurements along a transect across the lower epidermis revealed no correlation of stomatal aperture with the presence of major veins in the mesophyll tissue. Variation in guard-cell size and stomatal frequency on the lower leaf surface was also analysed. The guard cells were smallest and the stomata more frequent near to the leaf margins. The significance of the results is discussed in relation to measurements of leaf conductance and models of stomatal function.     

LeafAnalyser: a computational method for rapid and large-scale analyses of leaf shape variation

A comprehensive understanding of leaf shape is important in many investigations in plant biology. Techniques to assess variation in leaf shape are often time-consuming, labour-intensive and prohibited by complex calculation of large data sets. We have developed LeafAnalyser, software that uses image-processing techniques to greatly simplify the measurement of leaf shape variation. LeafAnalyser places a large number of evenly distributed landmarks along leaf margins and records the position of each automatically. We used LeafAnalyser to analyse the variation in 3000 leaves from 400 plants of Antirrhinum majus . We were able to summarise the major trends in leaf shape variation using a principal components (PC) analysis and assess the changes in size, width and tip-to-base asymmetry within our leaf library. We demonstrate how this information can be used to develop a model that describes the range and variation of leaf shape within standard wild-type lines, and illustrate the shape transformations that occur between leaf nodes. We also show that information from LeafAnalyser can be used to identify novel trends in shape variation, as low-variance PCs that only affect a subset of position landmarks. These results provide a high-throughput method to calculate leaf shape variation that allows a large number of leaves to be visualised in higher-dimensional phenotypic space. To illustrate the applicability of LeafAnalyser we also calculated the leaf shape variation in 300 leaves from Arabidopsis thaliana .     

LEAF SHAPE EVOLUTION IN THE SOUTH AFRICAN GENUS PELARGONIUM L' HÉR. (GERANIACEAE)

Leaf shapes reflect complex assemblages of shape-determining elements, yet evolutionary studies tend to treat leaf shape as a single attribute, for example cordate or linear. As with all complex structures, individual elements of a leaf could theoretically evolve independently and at different rates to the extent permitted by genetic and functional limitations. We examined relative evolutionary lability of shape-determining elements in the highly diverse South African plant genus Pelargonium (Geraniaceae). We used SIMMAP to calculate Bayesian posterior probabilities for ancestral states of leaf-shape characters for major nodes across multiple phylogenetic trees. Trees were derived from a Bayesian analysis of DNA sequence data from four partitions. We found that shape elements differed in rates of character-state transformations across the tree. Leaf base, apex, and overall outline had low rates. Transformations in venation occurred at slightly higher rates and were associated with shifts in venation among major clades. Leaf margin type and overall leaf size showed intermediate rates, whereas high rates were observed in the extent of lamina lobing and functional leaf size. The results indicate that suites of elements characteristic of the recently evolved xerophytic lineage, for example pinnate venation, dissected lamina, and entire margins, were acquired piecemeal over nested levels of the phylogeny.     

Regulation of Leaf Shape in the Solanifolia Mutant of Tomato (Lycopersicon esculentum) by Plant Growth Substances

The solanifolia mutant (sf/sf) of tomato (Lycopersicon esculentum)produces leaves consisting of leaflets with entire margins,unlike the lobed leaflets of normal plants. Normal plants treatedwith gibberellic acid (GA₃) produced leaves with entire marginswhereas mutant plants exposed to 2-chloroethyl-trimethyl ammoniumchloride (CCC)—an inhibitor of gibberellin biosynthesis—producedlobing of leaflets. The leaf area of the mutant was significantlygreater than that of the normal, but was not significantly differentfrom GA₃-treated normal leaves. Similarly, in CCC-treated mutantleaves the leaf area was not different from that of normal untreatedleaves. These observations suggest that the sf/sf mutation affectsthe leaf shape through its effect on endogenous gibberellinsand/or inhibitory substances. Leaf shape, Lycopersicon esculentum, plant growth substances, tomato     

Limiting the potential for intraspecific competition: regulation of Trioza eugeniae oviposition on unexpanded leaf tissue

1. Trioza eugeniae (Homoptera: Triozidae) females oviposit pre‐ ferentially on the margin of young, incompletely expanded Syzygium paniculatum leaves. As the density of eggs on the leaf increases, females will oviposit on the surface of the leaf blades. 2. The survivorship of eggs laid on the abaxial leaf midrib and near the abaxial leaf margin was significantly greater than that of eggs laid on the leaf margin. 3. Controlled environment studies demonstrated that maximum carrying capacity for T. eugeniae nymphs on a fully expanded leaf is approximated closely by the number of eggs that can be laid on the margin of a young, unexpanded leaf. 4. In field populations, nymphal densities fell below the hypothetical carrying capacity calculated from laboratory studies on 71% of the leaves examined. 5. The leaf margin appears to be a proximal cue used by ovipositing females for resources available to developing nymphs. Females that limit oviposition to available leaf margins appear to improve the probability of survival of their progeny through reduced intraspecific competition.     

Differences in Leaf Flammability,Leaf Traits and Flammability-Trait Relationships between Native and Exotic Plant Species of Dry Sclerophyll Forest

The flammability of plant leaves influences the spread of fire through vegetation. Exotic plants invading native vegetation may increase the spread of bushfires if their leaves are more flammable than native leaves. We compared fresh-leaf and dry-leaf flammability (time to ignition) between 52 native and 27 exotic plant species inhabiting dry sclerophyll forest. We found that mean time to ignition was significantly faster in dry exotic leaves than in dry native leaves. There was no significant native-exotic difference in mean time to ignition for fresh leaves. The significantly higher fresh-leaf water content that was found in exotics, lost in the conversion from a fresh to dry state, suggests that leaf water provides an important buffering effect that leads to equivalent mean time to ignition in fresh exotic and native leaves. Exotic leaves were also significantly wider, longer and broader in area with significantly higher specific leaf area–but not thicker–than native leaves. We examined scaling relationships between leaf flammability and leaf size (leaf width, length, area, specific leaf area and thickness). While exotics occupied the comparatively larger and more flammable end of the leaf size-flammability spectrum in general, leaf flammability was significantly correlated with all measures of leaf size except leaf thickness in both native and exotic species such that larger leaves were faster to ignite. Our findings for increased flammability linked with larger leaf size in exotics demonstrate that exotic plant species have the potential to increase the spread of bushfires in dry sclerophyll forest.     

Leaf anatomy of a rheophyte,Dendranthema yoshinaganthum (Asteraceae), and of hybrids between D. yoshinaganthumand a closely related non-rheophyte,D. indicum

The rheophyte Dendranthema yoshinaganthum(Makino ex Kitam.) Kitam. is endemic to a region along the Naka River in Shikoku, Japan. Resembling other rheophytes that have adapted to environments along rivers, D. yoshinaganthumhas narrow leaves. In the native habitat of D. yoshinaganthumin Tokushima, Japan, natural hybridization with Dendranthema indicum (L.) Des Moul. has resulted in a species complex with conspicuous variation in leaf shape and, in particular, in leaf width. In this study, D. yoshinaganthum and hybrids with natural variation in leaf shape were cultivated under identical conditions and the anatomy of their leaves was examined. Variation in leaf width in the species complex was attributable, for the most part, to variation in the number of cells per leaf lamina and also, to some extent, to variation in cell size. This is the first report of natural variation in leaf width that is attributable to variation in both the size and the number of leaf cells.     

从水力结构的角度解释生长温度对烟草叶片形状和大小的影响（英文）

叶片形状和大小在不同的生长温度下变化非常大,但少有从水力结构的角度解释其变化原因的研究。本研究测定了生长于两个不同温度下(24℃/18℃昼/夜;32℃/26℃昼/夜)的烟草叶片的解剖结构,导水率,叶片长宽比和叶面积。生长在24℃/18℃下的烟草叶片与生长在32℃/18℃的叶片相比,更狭窄,并有更小的叶柄导管直径,更低的叶脉密度和导水率。然而,在不同的生长温度下,烟草叶面积并没有显著差异。叶片导水率与叶脉密度呈正相关,但与叶片长宽比呈负相关。结果表明在不同的生长温度下叶片解剖结构和叶片导水率可能对于改变叶宽比起着重要作用。