Deciphering the evolutionary development of the "Chinese lantern" within Solanaceae.

MAIN CONCLUSION: The key genetic variation underlying the evo-devo of ICS in Solanaceae may be further pinpointed using an integrated strategy of forward and reverse genetics studies under the framework of phylogeny. The calyx of Physalis remains persistent throughout fruit development. Post-flowering, the fruiting calyx is inflated rapidly to encapsulate the berry, giving rise to a "Chinese lantern" structure called inflated calyx syndrome (ICS). It is unclear how this novelty arises. Over the past 2 decades, the role of MADS-box genes in the evolutionary development (evo-devo) of ICS has mainly been investigated within Solanaceae. In this review, we analyze the main achievements, challenges, and new progress. ICS acts as a source for fruit development, provides a microenvironment to protect fruit development, and assists in long-distance fruit dispersal. ICS is a typical post-floral trait, and the onset of its development is triggered by specific developmental signals that coincide with fertilization. These signals can be replaced by exogenous gibberellin and cytokinin application. MPF2-like heterotopic expression and MBP21-like loss have been proposed to be two essential evolutionary events for ICS origin, and manipulating the related MADS-box genes has been shown to affect the ICS size, sepal organ identity, and/or male fertility, but not completely disrupt ICS. Therefore, the core genes or key links in the ICS biosynthesis pathways may have undergone secondary mutations during evolution, or they have not yet been pinpointed. Recently, we have made some encouraging progress in acquiring lantern mutants in Physalis floridana. In addition to technological innovation, we propose an integrated strategy to further analyze the evo-devo mechanisms of ICS in Solanaceae using forward and reverse genetics studies under the framework of phylogeny.

Family ties: Root-root communication within Solanaceae.

Root-root communication effects on several physiological and metabolic aspects among Solanaceae relatives were studied. We examined cherry (C) and field (F) tomato (Solanum lycopersicum) and bell pepper (B) (Capsicum annuum), comprising three degrees of relatedness (DOR): high (H-DOR; CC, FF and BB), medium (M-DOR; CF) and low (L-DOR; CB and FB). Plants were grown in pairs of similar or different plants on a paper-based and non-destructive root growth system, namely, rhizoslides. Root growth, including the proliferation of fine roots, and respiration increased as the DOR decreased and were highest in paired L-DOR plants, as was shown for root respiration that increased by 63, 110 and 88 % for C, F, and B when grown with B, B and F, respectively. On the other hand, root exudates of L-DOR plants had significantly lower levels of total organic carbon and protein than those of H-DOR plants, indicating different root-root communication between individuals with different DOR. Our findings indicate, for the first time, that carbon allocation to root growth, exudation and respiration depends on the degree of genetic relatedness, and that the degree of relatedness between individual plants plays a key role in the root-root communication within Solanaceae.

Fields of flowers with few strikes: how oligolectic bees manage their foraging behavior on Calibrachoa elegans (Solanaceae).

In specialized plant-pollinator associations, partners may exhibit adaptive traits, which favor the maintenance of the interaction. The association between Calibrachoa elegans (Solanaceae) and its oligolectic bee pollinator, Hexantheda missionica (Colletidae), is mutualistic and forms a narrowly specialized pollination system. Flowers of C. elegans are pollinated exclusively by this bee species, and the bees restrict their pollen resources to this plant species. The pollen presentation schedules of C. elegans were evaluated at the population level to test the hypothesis that H. missionica females adjust their foraging behavior to the resource offering regime of C. elegans plants. For this, the number of new flowers and anthers opened per hour (as a proxy for pollen offering) was determined, and pollen advertisement was correlated with the frequency of flower visits during the day. Preferences of female bees for flowers of different stages were also investigated, and their efficiency as pollinators was evaluated. Pollen offering by C. elegans was found to be partitioned throughout the day through scattered flower openings. Females of H. missionica indeed adjusted their foraging activity to the most profitable periods of pollen availability. The females preferred new, pollen-rich flowers over old ones and gathered pollen and nectar selectively according to flower age. Such behaviors must optimize female bee foraging efficiency on flowers. Female bees set 93% of fruit after a single visit. These findings guarantee their importance as pollinators and the persistence of the specialized plant-pollinator association.

Understanding the relations between Solanaceae crops and their pollinators: a global meta-network.

Crop-pollinator interactions are essential for world food security. Studying crop pollination from a network approach allows identification of target pollinators for conservation and management, and gaps in our knowledge. Solanaceae represents the third highest ranked family based on economic value, and its production is highly improved by animal pollination. This study aimed to integrate global data on solanaceous crop pollination and analyse the interaction patterns using a meta-network approach. Our questions were: (i) how are interactions structured and what are the structuring roles of species; and (ii) what are the main gaps in our knowledge? Data were obtained through a systematic review of the main scientific databases. The network structure was described using connectivity and modularity calculations, and the role of species using centrality metrics. The 251 pollinator species reported were in seven orders, mainly Hymenoptera (84.9%). The generalists Bombus and Apis species were the most common pollinators. The meta-network was modular, and all modules mostly included bees. Most species were peripherals, around 12% were connectors, and there were no module hubs. Apis mellifera was the only network hub (supergeneralist). The most important pollinators are the most managed pollinators worldwide; however, many native species play a role in structuring the meta-network. Main gaps include species of importance to pepper pollination, lack of species-specific identification, and the need for more robust experimental studies evaluating the pollination efficiency of native, manageable bees.

Genome-Wide Analysis of MBF1 Family Genes in Five Solanaceous Plants and Functional Analysis of SlER24 in Salt Stress.

Multiprotein bridging factor 1 (MBF1) is an ancient family of transcription coactivators that play a crucial role in the response of plants to abiotic stress. In this study, we analyzed the genomic data of five Solanaceae plants and identified a total of 21 MBF1 genes. The expansion of MBF1a and MBF1b subfamilies was attributed to whole-genome duplication (WGD), and the expansion of the MBF1c subfamily occurred through transposed duplication (TRD). Collinearity analysis within Solanaceae species revealed collinearity between members of the MBF1a and MBF1b subfamilies, whereas the MBF1c subfamily showed relative independence. The gene expression of SlER24 was induced by sodium chloride (NaCl), polyethylene glycol (PEG), ABA (abscisic acid), and ethrel treatments, with the highest expression observed under NaCl treatment. The overexpression of SlER24 significantly enhanced the salt tolerance of tomato, and the functional deficiency of SlER24 decreased the tolerance of tomato to salt stress. SlER24 enhanced antioxidant enzyme activity to reduce the accumulation of reactive oxygen species (ROS) and alleviated plasma membrane damage under salt stress. SlER24 upregulated the expression levels of salt stress-related genes to enhance salt tolerance in tomato. In conclusion, this study provides basic information for the study of the MBF1 family of Solanaceae under abiotic stress, as well as a reference for the study of other plants.

Ptr1 evolved convergently with RPS2 and Mr5 to mediate recognition of AvrRpt2 in diverse solanaceous species.

The Ptr1 (Pseudomonas tomato race 1) locus in Solanum lycopersicoides confers resistance to strains of Pseudomonas syringae pv. tomato expressing AvrRpt2 and Ralstonia pseudosolanacearum expressing RipBN. Here we describe the identification and phylogenetic analysis of the Ptr1 gene. A single recombinant among 585 F2 plants segregating for the Ptr1 locus was discovered that narrowed the Ptr1 candidates to eight nucleotide-binding leucine-rich repeat protein (NLR)-encoding genes. From analysis of the gene models in the S. lycopersicoides genome sequence and RNA-Seq data, two of the eight genes emerged as the strongest candidates for Ptr1. One of these two candidates was found to encode Ptr1 based on its ability to mediate recognition of AvrRpt2 and RipBN when it was transiently expressed with these effectors in leaves of Nicotiana glutinosa. The ortholog of Ptr1 in tomato and in Solanum pennellii is a pseudogene. However, a functional Ptr1 ortholog exists in Nicotiana benthamiana and potato, and both mediate recognition of AvrRpt2 and RipBN. In apple and Arabidopsis, recognition of AvrRpt2 is mediated by the Mr5 and RPS2 proteins, respectively. Phylogenetic analysis places Ptr1 in a distinct clade compared with Mr5 and RPS2, and it therefore appears to have arisen by convergent evolution for recognition of AvrRpt2.

Survey of Sensitivity to Fatty Acid-Amino Acid Conjugates in the Solanaceae.

Plants perceive insect herbivores via a sophisticated surveillance system that detects a range of alarm signals, including herbivore-associated molecular patterns (HAMPs). Fatty acid-amino acid conjugates (FACs) are HAMPs present in oral secretions (OS) of lepidopteran larvae that induce defense responses in many plant species. In contrast to eggplant (Solanum melongena), tomato (S. lycopersicum) does not respond to FACs present in OS from Manduca sexta (Lepidoptera). Since both plants are found in the same genus, we tested whether loss of sensitivity to FACs in tomato may be a domestication effect. Using highly sensitive MAP kinase (MAPK) phosphorylation assays, we demonstrate that four wild tomato species and the closely related potato (S. tuberosum) do not respond to the FACs N-linolenoyl-L-glutamine and N-linolenoyl-L-glutamic acid, excluding a domestication effect. Among other genera within the Solanaceae, we found that bell pepper (Capsicum annuum) is responsive to FACs, while there is a differential responsiveness to FACs among tobacco (Nicotiana) species, ranging from strong responsiveness in N. benthamiana to no responsiveness in N. knightiana. The Petunia lineage is one of the oldest lineages within the Solanaceae and P. hybrida was responsive to FACs. Collectively, we demonstrate that plant responsiveness to FACs does not follow simple phylogenetic relationships in the family Solanaceae. Instead, sensitivity to FACs is a dynamic ancestral trait present in monocots and eudicots that was repeatedly lost during the evolution of Solanaceae species. Although tomato is insensitive to FACs, we found that other unidentified factors in M. sexta OS induce defenses in tomato.

Unravelling the structure and function of the petal appendages in the tribe Schwenckieae (Solanaceae).

The tribe Schwenckieae (Solanaceae) is characterised by the presence of appendages on the corolla, a diagnostic trait for the group. These appendages constitute a median distal projection of the three-lobed petal and occur in the genera Melananthus and Schwenckia but are absent in Heteranthia. We investigated the micromorphology and anatomical structure of the appendages and lateral petal lobes of Schwenckia americana (two varieties), S. angustifolia, S. curviflora and S. novaveneciana, and Melananthus fasciculatus. We also performed histochemical tests to determine if the appendages are involved in the production of volatiles, acting as a fragrance secretory structure (osmophore). The appendages have a uniseriate epidermis, whose cells store phenolics and lipids. The parenchyma is starch-rich just prior to anthesis in all species studied. The sensory test and anatomical analyses identified scent-secreting tissues, not only in the appendages, but also in the lateral petal lobes, whose cells are papillose with a sculptured surface. The α-naphthol p-phenylenediamine (NADI) reaction detected volatile (essential oils) compounds in S. americana var. americana and S. americana var. angustifolia. We demonstrated the secretory tissues and the production of lipids in the corolla appendages of Schwenckia and Melananthus, which indicate their osmogenic function and probable scent emission to attract pollinators.

Rapid customization of Solanaceae fruit crops for urban agriculture.

Cultivation of crops in urban environments might reduce the environmental impact of food production1-4. However, lack of available land in cities and a need for rapid crop cycling, to yield quickly and continuously, mean that so far only lettuce and related 'leafy green' vegetables are cultivated in urban farms5. New fruit varieties with architectures and yields suitable for urban farming have proven difficult to breed1,5. We identified a regulator of tomato stem length (SlER) and devised a trait-stacking strategy to combine mutations for condensed shoots, rapid flowering (SP5G) and precocious growth termination (SP). Application of our strategy using one-step CRISPR-Cas9 genome editing restructured vine-like tomato plants into compact, early yielding plants suitable for urban agriculture. Field data confirmed that yields were maintained, and we demonstrated cultivation in indoor farming systems. Targeting the same stem length regulator alone in groundcherry, another Solanaceae plant, also enabled engineering to a compact stature. Our approach can expand the repertoire of crops for urban agriculture.

Integrating historical biogeography and environmental niche evolution to understand the geographic distribution of Datureae.

PREMISE: The distributions of plant clades are shaped by abiotic and biotic factors as well as historical aspects such as center of origin. Dispersals between distant areas may lead to niche evolution when lineages are established in new environments. Alternatively, dispersing lineages may exhibit niche conservatism, moving between areas with similar environmental conditions. Here we test these contrasting hypotheses in the Datureae clade (Solanaceae). METHODS: We used maximum likelihood methods to estimate the ancestral range of Datureae along with the history of biogeographic events. We then characterized the niche of each taxon using climatic and soil variables and tested for shifts in environmental niche optima. Finally, we examined how these shifts relate to the niche breadth of taxa and clades within Datureae and the degree of overlap between them. RESULTS: Datureae originated in the Andes and subsequently expanded its range to North America and non-Andean regions of South America. The ancestral niche, and that of most Datura and Trompettia species, is dry, while Brugmansia species likely shifted toward a more mesic environment. Nonetheless, most Datureae present moderate to high overlap in niche breadth today. CONCLUSIONS: The expansion of Datureae into North America was associated with niche conservatism, with dispersal into similarly dry areas as occupied by the ancestral lineage. Subsequent niche evolution, including the apparent shift to a mesic niche in Brugmansia, diversified the range of habitats occupied by species in the tribe Datureae but also led to significant niche overlap among the three genera.

Four new species of Capsicum (Solanaceae) from the tropical Andes and an update on the phylogeny of the genus.

Four new species of Capsicum (Capsiceae, Solanaceae) from Andean tropical forests in South America are described. Capsicum benoistii Hunz. ex Barboza sp. nov. (incertae sedis) is endemic to a restricted area in south-central Ecuador and is most similar to the more widespread C. geminifolium (Dammer) Hunz. (Colombia, Ecuador, and Peru). Capsicum piuranum Barboza & S. Leiva sp. nov. (Andean clade) is found in northern Peru (Department Piura) and is morphologically most similar to C. caballeroi M. Nee of the Bolivian yungas (Departments Santa Cruz and Cochabamba) but closely related to C. geminifolium and C. lycianthoides Bitter. Capsicum longifolium Barboza & S. Leiva sp. nov. (Andean clade) occurs from northern Peru (Departments Amazonas, Cajamarca, and Piura) to southern Ecuador (Province Zamora-Chinchipe), and is morphologically most similar to C. dimorphum (Miers) Kuntze (Colombia, Ecuador, and Peru). Capsicum neei Barboza & X. Reyes sp. nov. (Bolivian clade) is endemic to southeastern Bolivia (Departments Chuquisaca and Santa Cruz) in the Boliviano-Tucumano Forest, is morphologically most similar to another Bolivian endemic species C. minutiflorum Rusby (Hunz.), and is closely related to C. caballeroi. Complete descriptions, illustrations, distributions and conservation assessments of all new species are given. Chromosome numbers for C. piuranum and C. longifolium are also provided. Three of the new species were included in a new phylogenetic analysis for Capsicum; their positions were strongly resolved within clades previously recognized in the genus.

Crescendo, diminuendo and subito of the trumpets: winds of change in the concerted evolution between flowers and pollinators in Salpichroa (Solanaceae).

'Gradual' vs 'punctuated' and 'unidirectional' (only lengthening) vs. 'bidirectional'(lengthenings and shortenings) modes of evolution are explanations that compete to explain adaptive changes of flower tube length in angiosperm. The nightshade genus Salpichroa Miers, with 21 species mostly growing in the tropical Andes of southern South America, has the opportune qualities of including nearly 15-fold inter-specific variation in corolla tube length, as well as one species that is a candidate for participating in evolutionary escalation with the longest-billed hummingbird, Ensifera ensifera. We reconstructed the phylogenetic relationships using five molecular markers, the two plastid markers trnD-trnT and trnL, and three nuclear markers, ITS and two COSII, and estimated divergence times of the genus in order to reconstruct the history of both corolla tube length and pollination mode (i.e. hummingbirds, moths or multiple). We used comparative methods to determine whether corolla tube elongation/shortening is associated with shifts in pollination mode and to test, modes and rates of corolla tube change. We found evidence of both lengthening and shortening of corolla tubes. Evolutionary rates are consistent with rapid corolla tube length transitions that are only partly associated with shifts in pollination mode. Though 'punctuated' evolution (i.e. large changes predominantly at speciation events) explained corolla changes in the whole genus, 'gradual' evolution (i.e. gradual changes during a coevolutionary race with the same pollinator) was a better explanation for the change in the long-flowered clade, mostly pollinated by hummingbirds.

Stepwise evolution of floral pigmentation predicted by biochemical pathway structure.

Developmental pathways play a major role in influencing the distribution of naturally occurring phenotypes. For example, pathway structure and regulation could make some phenotypes inaccessible or restrict the routes through which phenotypes evolve. In this study, we examine floral anthocyanin pigments across the Solanaceae family and test whether patterns of phenotypic variation are consistent with predicted constraints based on the structure of the flavonoid biosynthetic pathway. We find that anthocyanin evolution occurs in a stepwise manner whereby transitions between the production of red mono hydroxylated pelargonidin pigments and blue trihydroxylated delphinidin pigments first passes through an intermediate step of producing purple dihydroxylated cyanidin pigments. Although the transitions between these three pigment types differ in frequency, we infer that these shifts are often reversible, suggesting that the functionality of the underlying biochemical pathway is generally conserved. Furthermore, our study finds that some pigment combinations are never observed, pointing to additional constraints on naturally occurring phenotypes. Overall, our findings provide insights into how the structure of an angiosperm-wide biochemical pathway has shaped macroevolutionary variation in floral pigmentation.

Why are red flowers so rare? Testing the macroevolutionary causes of tippiness.

Traits that have arisen multiple times yet still remain rare present a curious paradox. A number of these rare traits show a distinct tippy pattern, where they appear widely dispersed across a phylogeny, are associated with short branches and differ between recently diverged sister species. This phylogenetic pattern has classically been attributed to the trait being an evolutionary dead end, where the trait arises due to some short-term evolutionary advantage, but it ultimately leads species to extinction. While the higher extinction rate associated with a dead end trait could produce such a tippy pattern, a similar pattern could appear if lineages with the trait speciated slower than other lineages, or if the trait was lost more often that it was gained. In this study, we quantify the degree of tippiness of red flowers in the tomato family, Solanaceae, and investigate the macroevolutionary processes that could explain the sparse phylogenetic distribution of this trait. Using a suite of metrics, we confirm that red-flowered lineages are significantly overdispersed across the tree and form smaller clades than expected under a null model. Next, we fit 22 alternative models using HiSSE (Hidden State Speciation and Extinction), which accommodates asymmetries in speciation, extinction and transition rates that depend on observed and unobserved (hidden) character states. Results of the model fitting indicated significant variation in diversification rates across the family, which is best explained by the inclusion of hidden states. Our best fitting model differs between the maximum clade credibility tree and when incorporating phylogenetic uncertainty, suggesting that the extreme tippiness and rarity of red Solanaceae flowers makes it difficult to distinguish among different underlying processes. However, both of the best models strongly support a bias towards the loss of red flowers. The best fitting HiSSE model when incorporating phylogenetic uncertainty lends some support to the hypothesis that lineages with red flowers exhibit reduced diversification rates due to elevated extinction rates. Future studies employing simulations or targeting population-level processes may allow us to determine whether red flowers in Solanaceae or other angiosperms clades are rare and tippy due to a combination of processes, or asymmetrical transitions alone.

Convergent evolution of floral shape tied to pollinator shifts in Iochrominae (Solanaceae).

Flower form is one of many floral features thought to be shaped by pollinator-mediated selection. Although the drivers of variation in flower shape have often been examined in microevolutionary studies, relatively few have tested the relationship between shape evolution and shifts in pollination system across clades. In the present study, we use morphometric approaches to quantify shape variation across the Andean clade Iochrominae and estimate the relationship between changes in shape and shifts in pollination system using phylogenetic comparative methods. We infer multiple shifts from an ancestral state of narrow, tubular flowers toward open, bowl-shaped, or campanulate flowers as well as one reversal to the tubular form. These transitions in flower shape are significantly correlated with changes in pollination system. Specifically, tubular forms tend to be hummingbird-pollinated and the open forms tend to be insect-pollinated, a pattern consistent with experimental work as well as classical floral syndromes. Nonetheless, our study provides one of the few empirical demonstrations of the relationship between flower shape and pollination system at a macroevolutionary scale.

A novel R3 MYB transcriptional repressor associated with the loss of floral pigmentation in Iochroma.

Losses of floral pigmentation represent one of the most common evolutionary transitions in flower color, yet the genetic basis for these changes has been elucidated in only a handful of cases. Here we used crossing studies, bulk-segregant RNA sequencing, phylogenetic analyses and functional tests to identify the gene(s) responsible for the transition to white flowers in Iochroma loxense. Crosses between I. loxense and its blue-flowered sister species, I. cyaneum, suggested that a single locus controls the flower color difference and that the white allele causes a nearly complete loss of pigmentation. Examining sequence variation across phenotypic pools from the crosses, we found that alleles at a novel R3 MYB transcription factor were tightly associated with flower color variation. This gene, which we term MYBL1, falls into a class of MYB transcriptional repressors and, accordingly, higher expression of this gene is associated with downregulation of multiple anthocyanin pigment pathway genes. We confirmed the repressive function of MYBL1 through stable transformation of Nicotiana. The mechanism underlying the evolution of white flowers in I. loxense differs from that uncovered in previous studies, pointing to multiple mechanisms for achieving fixed transitions in flower color intensity.

Epigenetic regulation of the expression of WRKY75 transcription factor in response to biotic and abiotic stresses in Solanaceae plants.

KEY MESSAGE: SlyWRKY75: gene expression was induced in response to biotic stresses, especially in Botrytis cinerea-infected tomato plants, in which Sly-miR1127-3p is a putative SlyWRKY75 regulator and epigenetic marks were detected. WRKY75 transcription factor involved in Pi homeostasis was recently found also induced in defense against necrotrophic pathogens. In this study, we analyzed by RT-qPCR the expression of SlyWRKY75 gene in tomato plants in response to abiotic stresses (drought or heat) and biotic stresses (Colorado potato beetle larvae infestation, Pseudomonas syringae or Botrytis cinerea infection) being only differentially expressed following biotic stresses, especially upon B. cinerea infection (55-fold induction). JA and JA-Ile levels were significantly increased in tomato plants under biotic stresses compared with control plants, indicating that SlyWRKY75 might be a transcriptional regulator of the JA pathway. The contribution of miRNAs and epigenetic molecular mechanisms to the regulation of this gene in B. cinerea-infected tomato plants was explored. We identified a putative Sly-miR1127-3p miRNA predicted to bind the intronic region of the SlyWRKY75 genomic sequence. Sly-miR1127-3p miRNA was repressed in infected plants (0.4-fold) supporting that it might act as an epigenetic regulation factor of SlyWRKY75 gene expression rather than via the post-transcriptional mechanisms of canonical miRNAs. It has been proposed that certain miRNAs can mediate DNA methylation in the plant nucleus broadening miRNA functions with transcriptional gene silencing by targeting intron-containing pre-mRNAs. Histone modifications analysis by chromatin immunoprecipitation (ChIP) demonstrated the presence of the activator histone modification H3K4me3 on SlyWRKY75 transcription start site and gene body. The induction of this gene in response to B. cinerea correlates with the presence of an activator mark. Thus, miRNAs and chromatin modifications might cooperate as epigenetic factors to modulate SlyWRKY75 gene expression.

A model for the irradiance responses of photosynthesis.

The analysis of the irradiance responses of photosynthetic processes, such as the quantum efficiencies of electron transport by photosystems I and II (PSI and PSII) or the rate of carbon dioxide fixation, is limited by the lack of mechanistically based analytical model for these processes. Starting with a model of P700 redox state, we develop a series of analytical functions which can be used to fit the irradiance responses of the quantum yields for electron transport by PSI and PSII, the irradiance responses of electron transport by PSI and PSII, and even the irradiance response of the fixation rate of carbon dioxide. These functions depend on two or three parameters so they can be fit to typical irradiance response data. We illustrate by example the use of these functions in various applications and discuss further use and development of the basic model described in detail here.

Self-compatibility is over-represented on islands.

Because establishing a new population often depends critically on finding mates, individuals capable of uniparental reproduction may have a colonization advantage. Accordingly, there should be an over-representation of colonizing species in which individuals can reproduce without a mate, particularly in isolated locales such as oceanic islands. Despite the intuitive appeal of this colonization filter hypothesis (known as Baker's law), more than six decades of analyses have yielded mixed findings. We assembled a dataset of island and mainland plant breeding systems, focusing on the presence or absence of self-incompatibility. Because this trait enforces outcrossing and is unlikely to re-evolve on short timescales if it is lost, breeding system is especially likely to reflect the colonization filter. We found significantly more self-compatible species on islands than mainlands across a sample of > 1500 species from three widely distributed flowering plant families (Asteraceae, Brassicaceae and Solanaceae). Overall, 66% of island species were self-compatible, compared with 41% of mainland species. Our results demonstrate that the presence or absence of self-incompatibility has strong explanatory power for plant geographical patterns. Island floras around the world thus reflect the role of a key reproductive trait in filtering potential colonizing species in these three plant families.

Differentiation in drought tolerance mirrors the geographic distributions of alpine plants on the Qinghai-Tibet Plateau and adjacent highlands.

Climatic tolerance, especially drought tolerance, is one of the major factors shaping the geographic distributions of plant species. Thus, the general decline in rainfall from the Himalaya-Hengduan Mountains (HHM) to the inner Qinghai-Tibet Plateau (QTP) might account for the significant differences in species distributions and richness between the two regions. To test this hypothesis, we conducted a water stress experiment using four Anisodus species (A. tanguticus, A. luridus, A. carniolicoides and A. acutangulus), which were treated with different levels of water stress in a glasshouse, and examined their differences in physiological responses. The results suggest that A. tanguticus, which inhabits the inner QTP, generally has higher fitness under severe water stress than the other species based on its high root:shoot ratio, long-term water use efficiency and photosynthetic rate, indicating that it possesses a genetically based drought tolerance mechanism. Our results suggest that plant species inhabiting the inner QTP may be more drought tolerant than those inhabiting the HHM regions. This provides a new example supporting the hypothesis that climatic tolerance plays a major role in shaping plant distributions on the QTP and its adjacent highlands and presents new insights into the patterns of geographic distribution and diversity of the plants inhabiting these areas.