Modulation of salt-induced stress impact in Gladiolus grandiflorus L. by exogenous application of salicylic acid.

Salinity is challenging threats to the agricultural system and leading cause of crop loss. Salicylic acid (SA) is an important endogenous signal molecule, which by regulating growth and physiological processes improves the plant ability to tolerate salt stress. Considering the prime importance of Gladiolus grandiflorus (L.) in the world's cut-flower market, the research work was undertaken to elucidate salinity tolerance in G. grandiflorus by exogenous application of SA irrigated with saline water. Results revealed that increasing salinity (EC: 2, 4 and 6 dS m-1) considerably altered morpho-growth indices (corm morphology and plant biomass) in plants through increasing key antioxidants including proline content and enzymes activity (superoxide dismutase, catalase and peroxidase), while negatively affected the total phenolic along with activity of defense-related enzymes (phenylalanine ammonia lyase, and polyphenol oxidase activity). SA application (50-200 ppm) in non-saline control or saline conditions improved morpho-physiological traits in concentration-dependent manners. In saline conditions, SA minimized salt-stress by enhancing chlorophyll content, accumulating organic osmolytes (glycine betaine and proline content), total phenolic, and boosting activity of antioxidant and defense-related enzymes. Principle component analysis based on all 16 morphological and physiological variables generated useful information regarding the classification of salt tolerant treatment according to their response to SA. These results suggest SA (100 or 150 ppm) could be used as an effective, economic, easily available and safe phenolic agent against salinity stress in G. grandiflorus.

Foliar epidermal anatomical characteristics of taxa of Iris subg. Scorpiris Spach (Iridaceae) from Turkey.

Iris L. is one of the important genus of family Iridaceae, consist of 56 taxa naturally occurred in Turkey. The similarities and variations in the subgenus overlapping the taxonomic positions of the species in the subgenera and needs anatomical assessment especially by microscopic techniques. In this study, the taxonomic significance of leaf anatomical characters of 10 Iris subgenus Scorpiris taxa were studied in detail and the relationship among these taxa were evaluated using microscopy techniques. Fresh leaf samples of species were fixed in 70% alcohol solution for anatomical observation under microscope. Eleven different micromorphological features were statistically analyzed to delimit the species in subgenus. Based on morphological and anatomical similarities, we studied relationships among; (1) ssp. turcica, ssp. caucasica, I. nezahatiae and I. pseudocaucasica; (2) correlation between ssp. turcica and ssp. caucasica; (3) association of I. galatica, I. persica, ssp. margaretiae and ssp. stenophylla with each other; (4) relationship between ssp. stenophylla and ssp. margaretiae; and (5) relevance between I. aucheri and I. peshmeniana. Moreover, the taxonomy of subgenus Scorpiris has been discussed in detail with novel and diagnostic features based on micromorphological physiognomies. We found that four species in this study are endemic to Turkey, while seven are critically endangered geophytes in the country. The leaf anatomical characteristics of 10 taxa were divided into three groups. Main aim of this research was to study the taxonomy of the complex subgenus Scorpiris through microscopic techniques.

Can short-billed nectar thieving sunbirds replace long-billed sunbird pollinators in transformed landscapes?

Pollinator specialisation through exploitation barriers (such as long floral tubes) does not necessarily mean a lack of pollination when the favoured pollinator is rare or absent. Theory predicts that suboptimal visitors will contribute to plant reproduction in the absence of the most effective pollinator. Here I address these questions with Chasmanthe floribunda a long-tubed plant species in the Cape Floristic Region, which is reliant on one species of pollinator, the long-billed Malachite Sunbird. In contrast to short-billed sunbirds, the Malachite Sunbird occurs in lower abundance or is absent in transformed landscapes. Short-billed sunbirds rob and thieve nectar from long-tubed flowers, but their potential contribution towards pollination is unknown. Experiments assessing seed set after single flower visits were performed to determine whether thieving short-billed sunbirds can act as substitute pollinators. To determine whether short-billed sunbirds reduce pollen limitation in transformed areas, pollen supplementation was done by hand and compared to natural fruit set. Short billed sunbirds are unable to act as substitute pollinators, and seed set is significantly lower in the flowers that they visited, compared to flowers visited by long-billed sunbirds. This is substantiated on a landscape scale, where fruit production in Chasmanthe floribunda could artificially be increased by 35% in transformed landscapes, but not so in natural areas. These findings have important consequences for the management and conservation of long-tubed bird-pollinated plant species that exist in recently transformed landscapes. The potential vulnerability of specialised plant species in transformed landscapes is highlighted.

Cadmium translocation by contractile roots differs from that in regular, non-contractile roots.

BACKGROUND AND AIMS: Contractile roots are known and studied mainly in connection with the process of shrinkage of their basal parts, which acts to pull the shoot of the plant deeper into the ground. Previous studies have shown that the specific structure of these roots results in more intensive water uptake at the base, which is in contrast to regular root types. The purpose of this study was to find out whether the basal parts of contractile roots are also more active in translocation of cadmium to the shoot. METHODS: Plants of the South African ornamental species Tritonia gladiolaris were cultivated in vitro for 2 months, at which point they possessed well-developed contractile roots. They were then transferred to Petri dishes with horizontally separated compartments of agar containing 50 µmol Cd(NO3)2 in the region of the root base or the root apex. Seedlings of 4-d-old maize (Zea mays) plants, which do not possess contractile roots, were also transferred to similar Petri dishes. The concentrations of Cd in the leaves of the plants were compared after 10 d of cultivation. Anatomical analyses of Tritonia roots were performed using appropriately stained freehand cross-sections. KEY RESULTS: The process of contraction required specific anatomical adaptation of the root base in Tritonia, with less lignified and less suberized tissues in comparison with the subapical part of the root. These unusual developmental characteristics were accompanied by more intensive translocation of Cd ions from the basal part of contractile roots to the leaves than from the apical-subapical root parts. The opposite effects were seen in the non-contractile roots of maize, with higher uptake and transport by the apical parts of the root and lower uptake and transport by the basal part. CONCLUSIONS: The specific characteristics of contractile roots may have a significant impact on the uptake of ions, including toxic metals from the soil surface layers. This may be important for plant nutrition, for example in the uptake of nutrients from upper soil layers, which are richer in humus in otherwise nutrient-poor soils, and also has implications for the uptake of surface-soil pollutants.

Intraspecific divergence and convergence of floral tube length in specialized pollination interactions.

Floral tubes are often thought to be a consequence of adaptive specialization towards pollinator morphology. We explore floral tube length evolution within Tritoniopsis revoluta (Iridaceae), a species with considerable geographical tube length variation. We ask whether tube lengths of T. revoluta populations are associated with pollinator proboscis lengths, whether floral divergence occurs in the presence of different pollinators and whether floral convergence occurs between distantly related populations pollinated by the same pollinator. Finally, we ask whether tube length evolution is directional. Shifts between morphologically different pollinators were always associated with shifts in floral morphology, even when populations were very closely related. Distantly related populations had similar tube lengths when they were pollinated by the same pollinator. Shifts in tube length tended to be from short to long, although reversals were not infrequent. After correcting for the population-level phylogeny, there was a strong positive, linear relationship between floral tube length and pollinator proboscis length, suggesting that plants are functionally specialized on different pollinators at different sites. However, because tube length evolution in this system can be a bidirectional process, specialization to the local pollinator fauna is unlikely to result in evolutionary or ecological dead-ends such as canalization or range limitation.

New insights into Trimezieae (Iridaceae) phylogeny: what do molecular data tell us?

BACKGROUND AND AIMS: The Neotropical tribe Trimezieae are taxonomically difficult. They are generally characterized by the absence of the features used to delimit their sister group Tigridieae. Delimiting the four genera that make up Trimezieae is also problematic. Previous family-level phylogenetic analyses have not examined the monophyly of the tribe or relationships within it. Reconstructing the phylogeny of Trimezieae will allow us to evaluate the status of the tribe and genera and to examine the suitability of characters traditionally used in their taxonomy. METHODS: Maximum parsimony and Bayesian phylogenetic analyses are presented for 37 species representing all four genera of Trimezieae. Analyses were based on nrITS sequences and a combined plastid dataset. Ancestral character state reconstructions were used to investigate the evolution of ten morphological characters previously considered taxonomically useful. KEY RESULTS: Analyses of nrITS and plastid datasets strongly support the monophyly of Trimezieae and recover four principal clades with varying levels of support; these clades do not correspond to the currently recognized genera. Relationships within the four clades are not consistently resolved, although the conflicting resolutions are not strongly supported in individual analyses. Ancestral character state reconstructions suggest considerable homoplasy, especially in the floral characters used to delimit Pseudotrimezia. CONCLUSIONS: The results strongly support recognition of Trimezieae as a tribe but suggest that both generic- and species-level taxonomy need revision. Further molecular analyses, with increased sampling of taxa and markers, are needed to support any revision. Such analyses will help determine the causes of discordance between the plastid and nuclear data and provide a framework for identifying potential morphological synapomorphies for infra-tribal groups. The results also suggest Trimezieae provide a promising model for evolutionary research.

Patterns of style polymorphism in five species of the South African genus Nivenia (Iridaceae).

BACKGROUND AND AIMS: Heterostylous plants have been characterized by the presence of two or three discrete morphs that differ in their sex organ position within populations. This polymorphism is widely distributed among the angiosperms, but detailed studies are limited to few taxonomic groups. Although a small representation, evolutionary meaningful variations of the heterostylous syndrome have been reported when precise measurements of the sexual whorls were taken. A thorough exploration of groups where heterostyly has been reported should offer new opportunities to further testing the evolutionary hypotheses explaining heterostyly. Here, the traits defining heterostyly were explored in half of the species in Nivenia, the only genus of Iridiaceae where heterostyly has been reported. METHODS: Detailed morphometric analysis of the flower sexual whorls and some traits considered as ancillary are supplied to determine for each population (a) the kind of stylar polymorphism, (b) the morph ratio and (c) the degree of reciprocity between sexual whorls. Also the rates of assortative (within morph) versus disassortative (between morphs) pollen transfer were estimated by analysing pollen loads on stigmas. The association between floral phenotypic integration and the reciprocity between sexual whorls was estimated; both characteristics have been quoted as dependent on the accuracy of the fit between pollinators and flowers and therefore related to the efficiency of pollen transfer. KEY RESULTS: Different types of polymorphism, differing in their degree of reciprocity, were found in Nivenia. Effective disassortative mating appears to be common, since (a) all dimorphic populations show equal morph-ratios (isoplethy), and (b) the pollen placed on the stigmas of each morph is likely to be coming from the other (complementary) morph. The most reciprocal populations of the heterostylous species have also the highest values of phenotypical integration. CONCLUSIONS: Stigma height dimorphism, as opposed to distyly, is proven for the first time in Nivenia. The presence of different types of polymorphism within the genus is consistent with hypotheses of the evolution of heterostyly. The role of the pollinators as the leading force of the transition seems to be apparent, since floral integration is related to reciprocity.

Evolution and coexistence of pollination ecotypes in an African Gladiolus (Iridaceae).

Pollinator-mediated selection has been suggested as a key driver of speciation in plants. We examined the potential role of hawkmoth pollinators in driving allopatric divergence and maintaining sympatric coexistence of morphotypes in the African iris Gladiolus longicollis. Floral tube length in this species varies from 35 mm to 130 mm across its geographic range and reflects the prevailing tongue lengths of local hawkmoth assemblages. The distribution of floral tube lengths is bimodal with two relatively discrete categories--long (about 90 mm) or short (about 50 mm)--that match the bimodal distribution of hawkmoth tongue lengths in eastern South Africa. At a contact site between these two floral morphs, we found few individuals of intermediate length, suggesting limited gene flow between morphs despite their interfertility. A difference in flowering phenology appears to be the main isolating barrier between morphs at this site. Long- and short-tubed morphs differed markedly in the chemical composition of their floral fragrance, a trait that could be used as a cue for morph-specific foraging by hawkmoths. Positive directional selection on tube length was found to occur in both morphs.

Radiation of pollination systems in the Iridaceae of sub-Saharan Africa.

BACKGROUND: Seventeen distinct pollination systems are known for genera of sub-Saharan African Iridaceae and recurrent shifts in pollination system have evolved in those with ten or more species. Pollination by long-tongued anthophorine bees foraging for nectar and coincidentally acquiring pollen on some part of their bodies is the inferred ancestral pollination strategy for most genera of the large subfamilies Iridoideae and Crocoideae and may be ancestral for the latter. Derived strategies include pollination by long-proboscid flies, large butterflies, night-flying hovering and settling moths, hopliine beetles and sunbirds. Bee pollination is diverse, with active pollen collection by female bees occurring in several genera, vibratile systems in a few and non-volatile oil as a reward in one species. Long-proboscid fly pollination, which is apparently restricted to southern Africa, includes four separate syndromes using different sets of flies and plant species in different parts of the subcontinent. Small numbers of species use bibionid flies, short-proboscid flies or wasps for their pollination; only about 2 % of species use multiple pollinators and can be described as generalists. SCOPE: Using pollination observations for 375 species and based on repeated patterns of floral attractants and rewards, we infer pollination mechanisms for an additional 610 species. Matching pollination system to phylogeny or what is known about species relationships based on shared derived features, we infer repeated shifts in pollination system in some genera, as frequently as one shift for every five or six species of southern African Babiana or Gladiolus. Specialized systems using pollinators of one pollination group, or even a single pollinator species are the rule in the family. Shifts in pollination system are more frequent in genera of Crocoideae that have bilaterally symmetric flowers and a perianth tube, features that promote adaptive radiation by facilitating precise shifts in pollen placement, in conjunction with changes in flower colour, scent and tube length. CONCLUSIONS: Diversity of pollination systems explains in part the huge species diversity of Iridaceae in sub-Saharan Africa, and permits species packing locally. Pollination shifts are, however, seen as playing a secondary role in speciation by promoting reproductive isolation in peripheral, ecologically distinct populations in areas of diverse topography, climate and soils. Pollination of Iridaceae in Eurasia and the New World, where the family is also well represented, is poorly studied but appears less diverse, although pollination by both pollen- and oil-collecting bees is frequent and bird pollination rare.

Botany: specialized bird perch aids cross-pollination.

Birds may hover over or perch on flowers when feeding on nectar, and this assists cross-pollination if they then visit other plants. Here we investigate the curious sterile inflorescence axis of the South African Cape endemic 'rat's tail' plant (Babiana ringens, Iridaceae), whose function--unlike in other bird-pollinated plants--is exclusively to provide a perch for foraging birds. We find that this structure promotes the plant's mating success by causing the malachite sunbird (Nectarinia famosa), its main pollinator, to adopt a position ideal for the cross-pollination of its unusual ground-level flowers.

Pollinator-mediated selection on flower-tube length in a hawkmoth-pollinated Gladiolus (Iridaceae).

Darwin's mechanistic model whereby selection favours plants with flower tubes that exceed the tongue length of the primary pollinator, was tested using unmanipulated plants of the hawkmoth-pollinated South African iris, Gladiolus longicollis. The study population was characterized by exceptionally large phenotypic variation in flower-tube length (range 56-129 mm). Directional selection on tube length was revealed by a significant positive relationship between this trait and both fruit and seed set. Selection was attributed to the effect of tube length on pollen receipt, as supplemental hand pollinations showed fruit and seed set in the population to be pollen limited. Indirect selection on tube length may also occur through the correlation of this trait with inflorescence height, although direct selection on the latter trait was significant only for seed set. The main pollinators at the study site were individuals of the large hawkmoth Agrius convolvuli that had tongue lengths of 85-135 mm. Other hawkmoths had tongues that were much too short to reach the nectar in G. longicollis flowers and seldom carried pollen of G. longicollis. Flowers with tubes shorter than the tongues of A. convolvuli are apparently not effectively pollinated because stigmas do not contact the moth's head effectively. This study demonstrates that selection may occur among plants with natural phenotypic variation in flower-tube length, and supports Darwin's model of pollinator-mediated selection.