Colonization in an abandoned East-Mediterranean vineyard

Abstract. Previous studies on secondary succession in abandoned agricultural land in the Mediterranean area were carried out by the chronosequence method, including data from different sites. A unique opportunity to study secondary succession arose from a situation in which different parts of one homogeneous East-Mediterranean vineyard were abandoned for 5, 8, 15 and 35 yr, and did not suffer from any disturbance subsequently. Most of the perennial species that colonized the abandoned vineyard were fleshy fruited species, which apparently were dispersed by birds from the surrounding maquis into the vineyard. These bird-dispersed species were the first to be established, and were the dominant plant group according to dispersal modes. The abandoned vine plants and their supporting columns provided the birds with perching and feeding sites, enhancing the arrival of bird-dispersed species regardless of their life forms. Under these conditions the most important attribute that affected vegetation dynamics was seed dispersal mode. Trees were among the first to colonize in the vineyard, implying that no facilitation was needed for their establishment. Annual plant species were the only species to disappear during succession. Almost all perennial species which had arrived persisted in the vineyard, and no replacement of perennial species was found. The rate of succession was rapid, as expressed by the short time (8–15 yr) needed for the stabilization of species composition, for growth to average height of late succession trees, and for reaching high cover of the invading perennial species in the abandoned vineyard. The secondary succession described above differs from that in the western Mediterranean by the absence of perennial species replacement and its rapid rate. The possible causes are discussed.     

Plant coloration undermines herbivorous insect camouflage

The main point of our hypothesis "coloration undermines camouflage" is that many color patterns in plants undermine the camouflage of invertebrate herbivores, especially insects, thus exposing them to predation and causing them to avoid plant organs with unsuitable coloration, to the benefit of the plants. This is a common case of "the enemy of my enemy is my friend" and a visual parallel of the chemical signals that plants emit to call wasps when attacked by caterpillars. Moreover, this is also a common natural version of the well-known case of industrial melanism, which illustrates the great importance of plant-based camouflage for herbivorous insects and can serve as an independent test for our hypothesis. We claim that the enormous variations in coloration of leaves, petioles and stems as well as of flowers and fruits undermine the camouflage of invertebrate herbivores, especially insects. We assume that the same principle might operate in certain animal-parasite interactions. Our hypothesis, however, does not contrast or exclude other previous or future explanations of specific types of plant coloration. Traits such as coloration that have more than one type of benefit may be selected for by several agents and evolve more rapidly than ones with a single type of advantage.     

Stability of pre- and post-fire spatial structure of pine trees in Aleppo pine forest

The study reported here describes for the first time the similarity between pre- and post-fire spatial patterns of the trees in a Mediterranean pine forest demonstrating that the pre-fire ancestor microsite is occupied also by the next generation. Although Aleppo pine Pinus halepensis Mill, is an obligatory post-fire seeder, it is adapted to regenerate in its pre-fire growing microsite. thus keeping suitable growing sites from generation to generation. We studied the effect of the dead burned adult pines on the density and size of their recruited saplings 2, 5, 11 and 20 yr after fire. A comparison of pine sapling density and size was made between the "near" zone (under the former effect of the burned canopy) and the'far'zone (beyond the former effect of the burned canopy).
In the site 2 yr after fire, seedling density was 56% higher in the "far" zone than in the'near'zone, but seedling .size was similar. However in the site 20 yr after fire, densities were similar in both zones, but the size was bigger by 89% in the "near" zone. Thus, population recruitment after fire seems to peak near the burned pine trees rather than at u distance from them, in contrast to Janzen's original'distance hypothesis' model suggested for undisturbed rainforest. Mere we present a new hypothetical model for the spatial pattern of post-fire regeneration of obligate seeder tree species forming open forests. It is proposed that in such trees the microsites which were kept by the burned adult trees, which are killed by the fire, are also the favorable regeneration microsite for the post-fire generation.     

Spatiotemporal dynamics of recruitment in Aleppo pine (Pinus halepensis Miller)

Spatial and temporal aspects of recruitment play a central role in plant population and community dynamics and have important basic and applied implications. Here we summarize and discuss the results from studies of spatiotemporal dynamics of recruitment stages (seeds-seedlings-saplings) in Aleppo pine (Pinus halepensis) throughout the species' native range. Seed release is induced either by fire (pyriscence) or by drying atmospheric conditions (xeriscence), each generating a distinct temporal pattern. In both cases most seeds travel relatively short distances (<30 m). Pyriscence is not expected to promote long-distance seed dispersal, but xeriscence is associated with relatively strong winds that can transport seeds over 1 km and more, as predicted by a new mechanistic dispersal model. In the absence of fire, seed survival increases with distance from adults because of higher competition with adults and sibs and higher seed predation near the seed sources. New data provide further evidence for such distance-dependent seed predation and also show that predation rates vary among habitats and are lowest in times of high seed abundance. The resulting recruitment patterns in the absence of fire are characterized by rapid spread and complex spatiotemporal dynamics that are fairly unpredictable and give rise to variable age structure. In contrast, the spatial pattern of the first post-fire generation is highly predictable because it explicitly replicates the spatial pattern of the pre-fire population. Unlike fire-free regeneration in which multiple factors operating at various stages are likely to be important, post-fire regeneration is governed by a fairly specific set of factors (the chemical properties of ash) operating during a specific stage (saplings) and at a specific location (the canopy projection of large burned pines). Post-fire forests are therefore even-aged and have predictable spatiotemporal dynamics. Most studies of Aleppo pine recruitment have focused on the seedling and sapling stages and on post-fire regeneration; we call for greater attention to the seed stage and to more extensive sampling of all recruitment stages in both space and time.     

Pollination of a core flowering shrub species in Mediterranean phrygana: variation in pollinator diversity, abundance and effectiveness in response to fire

Fire in Mediterranean-type ecosystems produces catastrophic changes in plant-pollinator systems; the recovery of which has been studied by comparing an unburnt mature forest habitat with that of an adjacent recently burnt area (eight years post-fire). The composition, visitation profiles, and effectiveness of the taxonomically diverse pollinator assemblages found on a core nectar providing species ( Satureja thymbra : Lamiaceae) were examined in these two contrasting habitats. S. thymbra in the freshly burnt area had low nectar standing crop and relatively less diverse bee community than an unburnt area which had twice the nectar standing crop and a higher bee diversity and abundance. Both sites supported bee assemblages dominated by the non-native bumblebee Bombus terrestris . Spatio-temporal heterogeneity of nectar standing crops and microclimatic conditions were sufficient to explain the form and magnitude of the diurnal foraging profiles at each site in relation to species specific foraging and flight abilities. B. terrestris, Apis mellifera and native solitary bees were the three primary guilds visiting S. thymbra and varied in the efficiency with which they delivered conspecific pollen grains to receptive stigmas. A pollinator effectiveness index for these three guilds was calculated based on floral visitation rates and pollen delivery efficiency and reflected the actual levels of effectiveness of each guild within and across the two habitat types. There was no overall inter-community difference in pollination effectiveness as the bee assemblages in both habitats were sufficient to produce maximum fruit set in S. thymbra, though the relative contribution of each guild varied intra-communally. Pollen limitation was not found to occur in either habitat.     

Unripe red fruits may be aposematic

The unripe fruits of certain species are red. Some of these species disperse their seeds by wind (Nerium oleander, Anabasis articulata), others by adhering to animals with their spines (Emex spinosa) or prickles (Hedysarum spinosissimum). Certainly neither type uses red coloration as advertisement to attract the seed dispersing agents. Fleshy-fruited species (Rhamnus alaternus, Rubus sanguineus and Pistacia sp.), which disperse their seeds via frugivores, change fruit color from green to red while still unripe and then to black or dark blue upon ripening. The red color does not seem to function primarily in dispersal (unless red fruits form advertisement flags when there are already black ripe fruits on the plant) because the red unripe fruits of these species are poisonous, spiny, or unpalatable. The unripe red fruits of Nerium oleander are very poisonous, those of Rhamnus alaternus and Anabasis articulata are moderately poisonous, those of Rubus sanguineus are very sour, those of Pistacia sp. contain unpalatable resin and those of Emex spinosa and Hedysarum spinosissimum are prickly. We propose that these unripe red fruits are aposematic, protecting them from herbivory before seed maturation.     

The type I membrane protein EFF-1 is essential for developmental cell fusion

Multinucleate cells are widespread in nature, yet the mechanism by which cells fuse their plasma membranes is poorly understood. To identify animal fusogens, we performed new screens for mutations that abolish cell fusion within tissues of C. elegans throughout development. We identified the gene eff-1, which is expressed as cells acquire fusion competence and encodes a novel integral membrane protein. EFF-1 sequence motifs suggest physicochemical actions that could cause adjacent bilayers to fuse. Mutations in the extracellular domain of EFF-1 completely block epithelial cell membrane fusion without affecting other perfusion events such as cell generation, patterning, differentiation, and adhesion. Thus, EFF-1 is a key component in the mechanism of cell fusion, a process essential to normal animal development.     

The effect of fire on flower visitation rate and fruit set in four core-species in east Mediterranean scrubland

The recovery of vegetation following fire has been studied intensively in Mediterranean-type ecosystems. Little attention, however, has been given to floral traits, and almost no data have been collected on the effects of fire on pollinator activity and fruit-set. This paper reports the effects of fire on flower visitation rates and the possibly related fruit-set. We compared visitation rates of the main pollinators on four plant core-species in burned and adjacent unburned areas. Measurements were performed at an unburned phrygana (scrub lands), and at a burned area (5–7 years post-fire). Bumble bees and solitary bees were the main taxa of visitors, while few honeybees were recorded. Solitary bees were almost absent from the burned area. Fruit-set was significantly higher in the unburned area for three out of the four plant species. The lower fruit-set in the burned area was possibly the result of low activity of solitary bees which are the main effective pollinators of the examined species. We hypothesize that the populations of the solitary bees were diminished or extirpated either directly by the fire, or indirectly by the scarcity of nectar in the early post-fire years due to dominance of young pine and Cistus spp. seedlings. The short foraging range of the solitary bees and their slow invasion rate into the burned area may explain our results.     

Cripto binds transforming growth factor beta (TGF-beta) and inhibits TGF-beta signaling

Cripto is a developmental oncoprotein and a member of the epidermal growth factor-Cripto, FRL-1, Cryptic family of extracellular signaling molecules. In addition to having essential functions during embryogenesis, Cripto is highly expressed in tumors and promotes tumorigenesis. During development, Cripto acts as an obligate coreceptor for transforming growth factor beta (TGF-beta) ligands, including nodals, growth and differentiation factor 1 (GDF1), and GDF3. As an oncogene, Cripto is thought to promote tumor growth via mechanisms including activation of mitogenic signaling pathways and antagonism of activin signaling. Here, we provide evidence supporting a novel mechanism in which Cripto inhibits the tumor suppressor function of TGF-beta. Cripto bound TGF-beta and reduced the association of TGF-beta with its type I receptor, TbetaRI. Consistent with its ability to block receptor assembly, Cripto suppressed TGF-beta signaling in multiple cell types and diminished the cytostatic effects of TGF-beta in mammary epithelial cells. Furthermore, targeted disruption of Cripto expression by use of small inhibitory RNA enhanced TGF-beta signaling, indicating that endogenous Cripto plays a role in restraining TGF-beta responses.     

DAP kinase and DRP-1 mediate membrane blebbing and the formation of autophagic vesicles during programmed cell death

Death-associated protein kinase (DAPk) and DAPk-related protein kinase (DRP)-1 proteins are Ca+2/calmodulin-regulated Ser/Thr death kinases whose precise roles in programmed cell death are still mostly unknown. In this study, we dissected the subcellular events in which these kinases are involved during cell death. Expression of each of these DAPk subfamily members in their activated forms triggered two major cytoplasmic events: membrane blebbing, characteristic of several types of cell death, and extensive autophagy, which is typical of autophagic (type II) programmed cell death. These two different cellular outcomes were totally independent of caspase activity. It was also found that dominant negative mutants of DAPk or DRP-1 reduced membrane blebbing during the p55/tumor necrosis factor receptor 1-induced type I apoptosis but did not prevent nuclear fragmentation. In addition, expression of the dominant negative mutant of DRP-1 or of DAPk antisense mRNA reduced autophagy induced by antiestrogens, amino acid starvation, or administration of interferon-gamma. Thus, both endogenous DAPk and DRP-1 possess rate-limiting functions in these two distinct cytoplasmic events. Finally, immunogold staining showed that DRP-1 is localized inside the autophagic vesicles, suggesting a direct involvement of this kinase in the process of autophagy.