Comparative plant development: the time of the leaf?

A key problem in developmental biology is understanding the origin of morphological innovations. Comparative studies in plants with different leaf morphologies indicate that the developmental pathway defined by KNOTTED1-type homeodomain proteins could be involved in generating different leaf forms. The differential expression of regulatory proteins has emerged as an important factor in driving morphological innovations in the plant kingdom--an idea that is well supported by quantitative trait locus analyses.     

Control of Leaf and Vein Development by Auxin

Leaves are the main photosynthetic organs of vascular plants and show considerable diversity in their geometries, ranging from simple spoonlike forms to complex shapes with individual leaflets, as in compound leaves. Leaf vascular tissues, which act as conduits of both nutrients and signaling information, are organized in networks of different architectures that usually mirror the surrounding leaf shape. Understanding the processes that endow leaves and vein networks with ordered and closely aligned shapes has captured the attention of biologists and mathematicians since antiquity. Recent work has suggested that the growth regulator auxin has a key role in both initiation and elaboration of final morphology of both leaves and vascular networks. A key feature of auxin action is the existence of feedback loops through which auxin regulates its own transport. These feedbacks may facilitate the iterative generation of basic modules that underlies morphogenesis of both leaves and vasculature.Leaf form and vascular patterns provide some of the most impressive examples of the complexity of biological shapes generated in nature. A common feature of the development of the leaf lamina and vein networks is the repeated use of basic modules. For example, the iterative emergence of marginal leaf-shape elements, such as serrations, lobes, and leaflets (Fig. 1A–D), and the arrangement of successive orders of branched veins result in different types of leaf geometries and vascular patterns, respectively. Intriguingly, there is also congruence of leaf shape and vein layouts, such that, at least superficially, the pattern of vasculature formation is well aligned with the final geometry of the leaf lamina. These observations raise the questions of (1) what are the specific signaling pathways that sculpt leaf shape and vascular patterns, (2) to what degree lamina growth and vascular development share common genetic control, and finally (3) how coordination between leaf and vascular development is achieved and impacts on generation of final leaf shape and vein arrangement. Over the past 15 years, genetic approaches have led to substantial increase in our understanding of leaf and vascular development, and have provided good evidence that regulated activity of the small indolic growth regulator auxin provides important spatial cues for both processes. Such roles of auxin in different facets of leaf and vascular development is the focus of our article.Open in a separate windowFigure 1.Axes of leaf asymmetry and diversity of leaf shape. (A) A simple, serrated leaf of the Columbia ecotype of Arabidopsis thaliana. The proximo–distal (P–D) and medio–lateral (M–L) axes are indicated in the image. The asterisk marks one marginal serration. (B) The lobed leaf of the Arabidopsis thaliana relative Arabidopsis lyrata. The asterisk depicts the position of one lobe. Lobes are deep serrations, so the definition of an outgrowth as a serration or lobe is somewhat arbitrary. (C) The dissected leaf of Cardamine hirsuta. The asterisk marks a lateral leaflet. Leaflets are clearly defined as distinct units of the same leaf, which connect with the rachis (R) via a structure called a petiolule (Pu). (D) The dissected leaf of the cultivated tomato. Tomato demonstrates additional orders of dissection with respect to Cardamine hirsuta leaf and produces both primary leaflets (black asterisk) and secondary leaflets (red asterisk). (E) Scanning electron micrograph of the shoot apex of tomato. The white asterisk marks a leaf primordium (1) initiating from the meristem. The adaxial (yellow) and abaxial (orange) domains are marked on the subsequent developing leaf (2). Tomato is a compound leaf plant where leaflets are formed from the leaf blade soon after leaf initiation (a developing leaflet is marked by an arrow in leaf 3). Images in panels A–D are leaf silhouettes. Scale bars: (A–D) 1 cm, (E) 100 µm.     

Autoregulation of RCO by Low-Affinity Binding Modulates Cytokinin Action and Shapes Leaf Diversity

1. Download : Download high-res image (79KB)
2. Download : Download full-size image

     

Weeds of change: Cardamine hirsuta as a new model system for studying dissected leaf development

Cardamine hirsuta, a small crucifer closely related to the model organism Arabidopsis thaliana, offers high genetic tractability and has emerged as a powerful system for studying the genetic basis for diversification of plant form. Contrary to A. thaliana, which has simple leaves, C. hirsuta produces dissected leaves divided into individual units called leaflets. Leaflet formation requires activity of Class I KNOTTED1-like homeodomain (KNOX) proteins, which also promote function of the shoot apical meristem (SAM). In C. hirsuta, KNOX genes are expressed in the leaves whereas in A. thaliana their expression is confined to the SAM, and differences in expression arise through cis-regulatory divergence of KNOX regulation. KNOX activity in C. hirsuta leaves delays the transition from proliferative growth to differentiation thus facilitating the generation of lateral growth axes that give rise to leaflets. These axes reflect the sequential generation of cell division foci across the leaf proximodistal axis in response to auxin activity maxima, which are generated by the PINFORMED1 (PIN1) auxin efflux carriers in a process that resembles organogenesis at the SAM. Delimitation of C. hirsuta leaflets also requires the activity of CUP SHAPED COTYLEDON (CUC) genes, which direct formation of organ boundaries at the SAM. These observations show how species-specific deployment of fundamental shoot development networks may have sculpted simple versus dissected leaf forms. These studies also illustrate how extending developmental genetic studies to morphologically divergent relatives of model organisms can greatly help elucidate the mechanisms underlying the evolution of form.     

Early phase changes by concurrent endurance and strength training

To compare regimens of concurrent strength and endurance training, 26 male basketball players were matched for stature, body composition, and physical activity level. Subjects completed different training programs for 7 weeks, 4 days per week. Groups were as follows: (a) the strength group (S; n = 7) did strength training; (b) the endurance group (E; n = 7) did endurance training; (c) the strength and endurance group (S + E; n = 7) combined strength and endurance training; and (d) the control group (C; n = 5) had no training. The S + E group showed greater gains in Vo(2)max than the E group did (12.9% vs. 6.8%), whereas the S group showed a decline (8.8%). Gains were noted in strength and vertical jump performance for the S + E and S groups. The S + E group had better posttraining anaerobic power than the S group did (6.2% vs. 2.9%). No strength, power, or anaerobic power gains were present for the E and C groups. We conclude that concurrent endurance and strength training is more effective in terms of improving athletic performance than are endurance and strength training apart.     

From genes to plants via meristems

The Society for Experimental Biology organised a "Plant Frontier" meeting, which was recently held at the University of Sheffield, UK. One of the sessions of this broad meeting was on plant meristems, which covered a range of topics, including stem cells, patterning, long distance signalling and epigenetic regulation of meristem development.