Simcha Lev-Yadun

Calmodulin binding to glutamate decarboxylase is required for regulation of glutamate and GABA metabolism and normal development in plants.

Glutamate decarboxylase (GAD) catalyzes the decarboxylation of glutamate to CO2 and gamma‐aminobutyrate (GABA). GAD is ubiquitous in prokaryotes and eukaryotes, but only plant GAD has been shown to bind calmodulin (CaM). Here, we assess the role of the GAD CaM‐binding domain in vivo. Transgenic tobacco plants expressing a mutant petunia GAD lacking the CaM‐binding domain (GADdeltaC plants) exhibit severe morphological abnormalities, such as short stems, in which cortex parenchyma cells fail to elongate, associated with extremely high GABA and low glutamate levels. The morphology of transgenic plants expressing the full‐length GAD (GAD plants) is indistinguishable from that of wild‐type (WT) plants. In WT and GAD plant extracts, GAD activity is inhibited by EGTA and by the CaM antagonist trifluoperazine, and is associated with a CaM‐containing protein complex of approximately 500 kDa. In contrast, GADdeltaC plants lack normal GAD complexes, and GAD activity in their extracts is not affected by EGTA and trifluoperazine. We conclude that CaM binding to GAD is essential for the regulation of GABA and glutamate metabolism, and that regulation of GAD activity is necessary for normal plant development. This study is the first to demonstrate an in vivo function for CaM binding to a target protein in plants.

Unravelling the evolution of autumn colours: an interdisciplinary approach

Leaf colour change is commonly observed in temperate deciduous forests in autumn. This is not simply a side effect of leaf senescence, and, in the past decade, several hypotheses have emerged to explain the evolution of autumn colours. Yet a lack of crosstalk between plant physiologists and evolutionary ecologists has resulted in slow progress, and so the adaptive value of this colour change remains a mystery. Here we provide an interdisciplinary summary of the current body of knowledge on autumn colours, and discuss unresolved issues and future avenues of research that might help reveal the evolutionary meaning of this spectacle of nature.

Differentiation of the ray system in woody plants

The regulation of vascular ray differentiation has received limited attention, despite the fact that vascular rays constitute an important part of the secondary body of plants. In this paper we review developmental aspects of the ray system and suggest a general hypothesis for the regulation of ray differentiation and evolution. In studies of ray differentiation, two basic factors should be taken into consideration: 1) the normal gradual increase in ray size in relation to age, distance from the pith, and distance from the young leaves; and 2) the influence of wound effects on the size, structure, and spacing of rays. The relationships between the rate of cambial activity and secondary xylem differentiation are not clearly understood. There are contrasting results on the relationships between ray number and rate of radial growth. The rate of radial growth (= rate of cambial activity) is not the regulating mechanism of ray characteristics. Bünning (1952, 1965) proposed that rays are distributed regularly in the tissue, as the outcome of an inhibitory influence expressed by them. However, Bünning’s hypothesis contradicts a basic feature of the vascular ray system, namely, fusion of rays. Detailed histological studies of the secondary xylem revealed that proximity to and contact with rays plays a major role in the survival of fusiform initials in the cambium (Bannan, 1951, 1953). Such evidence led Ziegler (1964) to suggest that since the cambium is supplied predominantly via the rays, this is an effective feedback regulative system for an equidistant arrangement of the rays. The hypothesis that rays are induced and controlled by a radial signal flow seems to be the best explanation for the structure and spacing of rays. The formation of a polycentric ray—a special case of “ray” initiation inside a vascular ray—supports the idea that radial signal flow occurs within the rays (Lev-Yadun & Aloni, 1991a). This idea is also supported by findings fromQuercus species in which aggregate rays in the xylem disperse naturally in branch junctions and, following partial girdling, leave a longitudinal narrow bridge of cambium and bark as a result of enhanced axial signal flow (of auxin and other growth regulators) (Lev-Yadun & Aloni, 1991b). The longitudinally elongated shape of rays is their response to axial signal flows (mainly the polar auxin flow). Two methods have been used to study the evolution of the ray system: 1) statistical studies of the relationships between vessel and ray characteristics in many species, when vessel characteristics were the evolutionary standard, and 2) comparison of ray characteristics in fossils originating from several geological eras. We suggest that evolution of the ray system reflects changes in the relations between radial and axial signal flows.

Cambial Activity of Evergreen and Seasonal Dimorphics Around the Mediterranean

The annual rhythm of cambial activity in various Mediterranean evergreens and seasonal dimorphics is compared on the basis of a literature survey. Two main rhythms of activity can be distinguished:

What Do Red and Yellow Autumn Leaves Signal

The widespread phenomenon of red and yellow autumn leaves has recently attracted considerable scientific attention. The fact that this phenomenon is so prominent in the cooler, temperate regions and less common in warmer climates is a good indication of a climate-specific effect. In addition to the putative multifarious physiological benefits, such as protection from photoinhibition and photo-oxidation, several plant/animal interaction functions for such coloration have been proposed. These include (1) that the bright leaf colors may signal frugivores about ripe fruits (fruit flags) to enhance seed dispersal; (2) that they signal aphids that the trees are well defended (a case of Zahavi’s handicap principle operating in plants); (3) that the coloration undermines herbivore insect camouflage; (4) that they function according to the “defense indication hypothesis,” which states that red leaves are chemically defended because anthocyanins correlate with various defensive compounds; or (5) that because sexual reproduction advances the onset of leaf senescence, the pigments might indicate to sucking herbivores that the leaves have low amounts of resources. Although the authors of hypotheses 3, 4, and 5 did not say that bright autumn leaves are aposematic, since such leaves are chemically defended, unpalatable, or both, we suggest that they are indeed aposematic. We propose that in addition to the above-mentioned hypotheses, autumn colors signal to herbivorous insects about another defensive plant property: the reliable, honest, and critical information that the leaves are about to be shed and may thus cause their mortality. We emphasize that all types of defensive and physiological functions of autumn leaves may operate simultaneously.

The Salt-Stress Signal Transduction Pathway That Activates the gpx1 Promoter Is Mediated by Intracellular H2O2, Different from the Pathway Induced by Extracellular H2O2

Several genes encoding putative glutathione peroxidase have been isolated from a variety of plants, all of which show the highest homology to the phospholipid hydroperoxide isoform. Several observations suggest that the proteins are involved in biotic and abiotic stress responses. Previous studies on the regulation of gpx1, the Citrus sinensis gene encoding phospholipid hydroperoxide isoform, led to the conclusion that salt-induced expression of gpx1 transcript and its encoded protein is mediated by oxidative stress. In this paper, we describe the induction of gpx1 promoter:uidA fusions in stable transformants of tobacco (Nicotiana tabacum) cultured cells and plants. We show that the induction of gpx1 by salt and oxidative stress occurs at the transcriptional level. gpx1 promoter analysis confirmed our previous assumption that the salt signal is transduced via oxidative stress. We used induction of the fusion construct to achieve better insight into, and to monitor salt-induced oxidative stress. The gpx1 promoter responded preferentially to oxidative stress in the form of hydrogen peroxide, rather than to superoxide-generating agents. Antioxidants abolished the salt-induced expression of gpx1 promoter, but were unable to eliminate the induction by H2O2. The commonly employed NADPH-oxidase inhibitor diphenyleneiodonium chloride and catalase inhibited the H2O2-induced expression of gpx1 promoter, but did not affect its induction by salt. Our results led us to conclude that salt induces oxidative stress in the form of H2O2, its production occurs in the intracellular space, and its signal transduction pathway activating the gpx1 promoter is different from the pathway induced by extracellular H2O2.

Agricultural Origins: Centers and Noncenters; A Near Eastern Reappraisal

Understanding the evolutionary history of crop plants is fundamental to our understanding of their respective adaptation profiles, which in turn, is a key element in securing future yield and quality improvement. Central topics in this field concern the mono- or polyphyletic origin of crop plants, and our ability to identify the geographic location where certain crop plants have originated. Understanding the geographical pattern of domestication may also assist in reconstructing the cultural processes underlying the Neolithic (agricultural) Revolution. Here we review prevailing views on the geographic pattern of Near Eastern plant domestication, and highlight the distinction between genetic domestication events and independent cultural events. A critical evaluation of the wealth of newly published geobotanical, genetic, and archaeological data provides strong support in favor of a specific core area in southeastern Turkey where most, if not all, founder Near Eastern crops were likely domesticated.

The Chickpea, Summer Cropping, and a New Model for Pulse Domestication in the Ancient Near East

The widely accepted models describing the emergence of domesticated grain crops from their wild type ancestors are mostly based upon selection (conscious or unconscious) of major features related either to seed dispersal (nonbrittle ear, indehiscent pod) or free germination (nondormant seeds, soft seed coat). Based on the breeding systems (self‐pollination) and dominance relations between the allelomorphs of seed dispersal mode and seed dormancy, it was postulated that establishment of the domesticated forms and replacement of the wild ancestral populations occurred in the Near East within a relatively short time. Chickpea (Cicer arietinum L.), however, appears as an exception among all other “founder crops” of Old World agriculture because of its ancient conversion into a summer crop. The chickpea is also exceptional because its major domestication trait appears to be vernalization insensitivity rather than pod indehiscence or free germination. Moreover, the genetic basis of vernalization response in wild chickpea (Cicer reticulatum Ladiz.) is polygenic, suggesting that a long domestication process was imperative due to the elusive phenotype of vernalization nonresponsiveness. There is also a gap in chickpea remains in the archaeological record between the Late Prepottery Neolithic and the Early Bronze Age. Contrary to the common view that Levantine summer cropping was introduced relatively late (Early Bronze Age), we argue for an earlier (Neolithic) Levantine origin of summer cropping because chickpea, when grown as a common winter crop, was vulnerable to the devastating pathogen Didymella rabiei, the causal agent of Ascochyta blight. The ancient (Neolithic) conversion of chickpea into a summer crop required seasonal differentiation of agronomic operation from the early phases of the Neolithic revolution. This topic is difficult to deal with, as direct data on seasonality in prehistoric Old World field crop husbandry are practically nonexistent. Consequently, this issue was hardly dealt with in the literature. Information on the seasonality of ancient (Neolithic, Chalcolithic, and Early Bronze Age, calibrated 11,500 to 4,500 years before present) Near Eastern agriculture may improve our understanding of the proficiency of early farmers. This in turn may provide a better insight into Neolithic agrotechniques and scheduling. It is difficult to fully understand chickpea domestication without a Neolithic seasonal differentiation of agronomic practice because the rapid establishment of the successful Near Eastern crop package which included wheats, barley, pea, lentil, vetches, and flax, would have preempted the later domestication of this rare wild legume.

Plant Fiber Formation: State of the Art, Recent and Expected Progress, and Open Questions

Plant fibers are one of the most important renewable resources, used as raw material in the paper industry, and for various textiles and for composites. Fibers are structural components in timber and an energy-rich component of fuel-wood. For the plant itself, fibers are important in establishing plant architecture, as a source of mechanical support, in defence from herbivory, and in some cases as elements with contractile properties, resembling those of muscles. In addition, fibers may store ergastic carbon resources and water. Here, we review various aspects of fiber development such as initiation, elongation, cell wall formation and multinuclearity, discuss open questions and propose directions for further research. Most of the recent progress in fiber formation biology, especially in cell wall structure and chemistry, emerged from studies of only a few model plants including flax, Populus spp., Eucalyptus spp., Arabidopsis thaliana and hemp. Considering the enormous importance of fibers to humanity, it is surprising how little is known about the biology of fiber formation.

Why do some thorny plants resemble green zebras

The rosette and cauline leaves of the highly thorny winter annual plant species of the Asteraceae in Israel (Silybum marianum) resemble green zebras. The widths of typical variegation bands were measured and found to be highly correlated with leaf length, length of the longest spines at leaf margins and the number of spines along leaf circumference. Thus, there is a significant correlation between the spinyness and strength of variegation. I propose that this is a special case of aposematic (warning) coloration.