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1.  Relative importance of phenotypic trait matching and species' abundances in determining plant–avian seed dispersal interactions in a small insular community 
AoB Plants  2015;7:plv017.
In this paper the authors take advantage of the simplicity of an insular community to evaluate the relative importance of species' phenotypic traits and species' abundance in determining fruit-avian disperser interactions, at both network and pairwise interaction levels. The authors innovatively include fruit nutrient compounds in fruit-avian network analyses. Although the best way to predict plant-avian interactions was based on both phenotypic traits and species abundance, the most important factor to explain these mutualistic interactions was fruit-beak size overlap, followed by species abundance and fruit nutrient compounds. This work will encourage further studies to look for similar patterns in more species-rich communities.
Network theory has provided a general way to understand mutualistic plant–animal interactions at the community level. However, the mechanisms responsible for interaction patterns remain controversial. In this study we use a combination of statistical models and probability matrices to evaluate the relative importance of species morphological and nutritional (phenotypic) traits and species abundance in determining interactions between fleshy-fruited plants and birds that disperse their seeds. The models included variables associated with species abundance, a suite of variables associated with phenotypic traits (fruit diameter, bird bill width, fruit nutrient compounds), and the species identity of the avian disperser. Results show that both phenotypic traits and species abundance are important determinants of pairwise interactions. However, when considered separately, fruit diameter and bill width were more important in determining seed dispersal interactions. The effect of fruit compounds was less substantial and only important when considered together with abundance-related variables and/or the factor ‘animal species’.
doi:10.1093/aobpla/plv017
PMCID: PMC4372831  PMID: 25750409
Dispersal; frugivory; mutualistic networks; oceanic islands; probability matrices
2.  Ecotypic variation in growth responses to simulated herbivory: trade-off between maximum relative growth rate and tolerance to defoliation in an annual plant 
AoB Plants  2015;7:plv015.
How fast plant genotypes grow before damage by herbivores has been theorized to impact negatively on the tolerance response to defoliation. Using a growth analytical approach in two ecotypes of Datura stramonium that differ in relative growth rate (before defoliation) and tolerance to defoliation, this study shows that slow growing plant genotypes exhibit the highest compensation (more trait expression) after defoliation, not only in relative growth rate, but also in fitness (number of seeds). At the intra-specific scale, a trade-off between the ability to grow under benign environmental conditions and the ability to tolerate resource limitation due to defoliation was detected.
It has been hypothesized that slow-growing plants are more likely to maximize above-ground biomass and fitness when defoliated by herbivores than those with an already high relative growth rate (RGR). Some populations of the annual herb Datura stramonium L. can tolerate foliar damage better than others. The physiological basis of this difference is examined here in a comparative study of two ecotypes that differ in tolerance and maximum growth rate, using a growth analytical approach. One hundred and fifty-four plants of each ecotype grown under controlled conditions were suddenly defoliated (35 % of total leaf area removed) and a similar sample size of plants remained undefoliated (control). Ontogenetic plastic changes in RGR and its growth components [net assimilation rate (NAR), specific leaf area and leaf weight ratio (LWR)] after defoliation were measured to determine whether these plastic changes maximize plant growth and fitness. Different ontogenetic phases of the response were discerned and increased RGR of defoliated plants was detected at the end of the experimental period, but brought about by a different growth component (NAR or LWR) in each ecotype. These changes in RGR are putatively related to increases in fitness in defoliated environments. At the intra-specific scale, data showed a trade-off between the ability to grow under benign environmental conditions and the ability to tolerate resource limitation due to defoliation.
doi:10.1093/aobpla/plv015
PMCID: PMC4372832  PMID: 25725085
Datura stramonium; defoliation tolerance; growth response coefficients; net assimilation rate; ontogenetic plasticity response to defoliation; plant relative growth rate
3.  Rainwater in cupulate bracts repels seed herbivores in a bumblebee-pollinated subalpine flower 
AoB Plants  2015;7:plv019.
The discovery of new adaptations of organisms to various environments always reminds us how little we know about nature. For example, rainwater surrounding sexual organs has been noted in several flowering plant groups. Sun and Huang used a bumblebee-pollinated alpine flower, Pedicularis rex (Orobanchaceae), which has cup-like bracts holding rainwater, as an experimental model to examine whether the water-filled cupulate bracts function to deter nectar robbers and/or seed herbivores. They found that neither nectar robbers nor legitimate pollinators discriminated water-drained flowers, but seed predation significantly increased in drained flowers, suggesting that water-filled bracts help protect the flowers from seed herbivores.
Floral herbivory may have deleterious effects on the reproductive success of flowering plants. However, plants may evolve floral traits that allow them to defend against herbivory in particular conditions. A bumblebee-pollinated subalpine herb, Pedicularis rex (Orobanchaceae), endemic to southwest China, has cup-like bracts that fill with rainwater, which submerges its corolla tubes. We hypothesized that these water-filled cupulate bracts function to deter nectar robbers and/or seed herbivores. To test these hypotheses, we experimentally drained bracts and measured both the response of mutualistic floral visitors and antagonistic nectar robbers and seed predators and their effects on seed production. Our observations revealed that neither nectar robbers nor legitimate pollinators discriminated between water-drained flowers and intact controls. However, seed predation significantly increased in drained flowers, suggesting that water-filled bracts help protect the flowers from seed herbivores. The water-filled bracts in P. rex may represent an adaptation to reduce floral herbivory in a high-rainfall environment.
doi:10.1093/aobpla/plv019
PMCID: PMC4392828  PMID: 25762460
Cupulate bract; floral herbivory; legitimate pollinator; nectar robber; Pedicularis rex; seed predation
4.  Germination and ultrastructural studies of seeds produced by a fast-growing, drought-resistant tree: implications for its domestication and seed storage 
AoB Plants  2015;7:plv016.
The inner layers of the seed coat that remain attached to the cotyledons probably play a role in seed dormancy of Moringa oleifera. Cotyledons of seeds stored for one year showed no sign of deterioration. In some cells of the three-year-old cotyledons, the membranes of the protein bodies were deteriorated. Cell deterioration was also marked by the collapse of the cell wall adjacent to the intercellular cavity. The decrease in seed viability during storage is associated with the loss in membrane integrity as confirmed by the increase in electrolyte leakage. The longevity of seeds can be extended if they are stored within their fruits under favourable conditions.
Seed ageing during storage is one of the main causes of reduction in seed quality and this results in loss of vigour and failure to thrive. Finding appropriate storage conditions to ameliorate deterioration due to ageing is, therefore, essential. Ultrastructural changes in cellular organelles during storage and seed germination rates are valuable indices of damage that occurs during seed ageing. There is increasing interest in Moringa oleifera Lam. because of its multiple uses as an agroforestry crop. Seeds of this species lose their viability within 6–12 months of harvest but no scientific information is available on the longevity of seed stored in the fruit (capsules). In most undeveloped countries, seeds are still stored inside the fruit by traditional methods in special handmade structures. In this experiment we tried to simulate these traditional storage conditions. Capsules of Moringa were stored at ambient room temperature for 12, 24 and 36 months. The ultrastructure, solute leakage and viability of seed were investigated. The ultrastructure of 1-year-old seed showed no sign of deterioration. It was evident, however, that some cells of the 3-year-old seed had deteriorated. The remnants of the outer and inner two integuments that remain tightly attached to the cotyledons probably play a role in seed dormancy. No significant difference was found between germination percentage of fresh and 1-year-old seed. The germination percentage decreased significantly from 2 years of storage onward. The decrease in seed viability during storage was associated with a loss in membrane integrity which was evidenced by an increase in electrolyte leakage. Our findings indicate that the longevity of M. oleifera seeds can be maintained if they are stored within their capsules.
doi:10.1093/aobpla/plv016
PMCID: PMC4379585  PMID: 25725083
Deterioration; dormancy; lipid bodies; membrane leakage; protein bodies; seed; storage
5.  Changes in plant species composition of coastal dune habitats over a 20-year period 
AoB Plants  2015;7:plv018.
Coastal sandy ecosystems are increasingly being threatened by human pressure, causing loss of biodiversity and habitat degradation. Using phytosociological relevés we conducted a re-visitation study in order to analyse changes in floristic composition during the last twenty years along the central Adriatic coast. We observed a significant increase in cover of fore dune and thermophilic species. Even though human activities are major driving forces of change in coastal dune vegetation, the species' cover increase may also be due to a moderate increment in average yearly temperature over the last two decades.
Coastal sandy ecosystems are increasingly being threatened by human pressure, causing loss of biodiversity, habitat degradation and landscape modifications. However, there are still very few detailed studies focussing on compositional changes in coastal dune plant communities over time. In this work, we investigated how coastal dune European Union (EU) habitats (from pioneer annual beach communities to Mediterranean scrubs on the landward fixed dunes) have changed during the last 20 years. Using phytosociological relevés conducted in 1989–90 and in 2010–12, we investigated changes in floristic composition over time. We then compared plant cover and the proportion of ruderal, alien and habitat diagnostic species (‘focal species’) in the two periods. Finally, we used Ellenberg indicator values to define the ‘preferences’ of the plant species for temperature and moisture. We found that only fore dune habitats showed significant differences in species cover between the two time periods, with higher plant cover in the more recent relevés and a significant increase in thermophilic species. Although previous studies have demonstrated consistent habitat loss in this area, we observed that all coastal dune plant communities remain well represented, after a 20-year period. However, fore dunes have been experiencing significant compositional changes. Although we cannot confirm whether the observed changes are strictly related to climatic changes, to human pressure or to both, we hypothesize that a moderate increment in average yearly temperature may have promoted the increase in plant cover and the spread of thermophilic species. Thus, even though human activities are major driving forces of change in coastal dune vegetation, at the community scale climatic factors may also play important roles. Our study draws on re-visitation studies which appear to constitute a powerful tool for the assessment of the conservation status of EU habitats.
doi:10.1093/aobpla/plv018
PMCID: PMC4398890  PMID: 25750408
Coastal dune zonation; diachronic analysis; phytosociological relevés; re-visitation study; vegetation changes
6.  Linkages of plant–soil feedbacks and underlying invasion mechanisms 
AoB Plants  2015;7:plv022.
Plant-soil feedbacks including soil pathogens, mycorrhizal costs/benefits and nutrient dynamics impact invasion directly or through interactions with other ecological processes. Here we describe how the mechanisms by which soil community processes influence plant growth overlap with several known and well-described mechanisms of plant invasion. We need to focus on the underlying mechanisms of soil feedbacks, and how they link to invasion along temporal scale.
Soil microbial communities and processes have repeatedly been shown to impact plant community assembly and population growth. Soil-driven effects may be particularly pronounced with the introduction of plants to non-native ranges, as introduced plants are not typically accompanied by transference of local soil communities. Here we describe how the mechanisms by which soil community processes influence plant growth overlap with several known and well-described mechanisms of plant invasion. Critically, a given soil community process may either facilitate or limit invasion, depending upon local conditions and the specific mechanisms of soil processes involved. Additionally, as soil communities typically consist of species with short generation times, the net consequences of plant–soil feedbacks for invasion trajectories are likely to change over time, as ecological and evolutionary adjustments occur. Here we provide an overview of the ecological linkages of plant–soil feedbacks and underlying mechanisms of invasion.
doi:10.1093/aobpla/plv022
PMCID: PMC4404623  PMID: 25784668
Invasion; native species; non-native ranges; soil communities; virulent pathogens
7.  Introduction to the Special Issue: Halophytes in a changing world 
AoB Plants  2015;7:plv020.
Climate change will bring about rising sea levels and increasing drought, both of which will contribute to increasing salinization in many regions of the world. There will be consequent effects on our crops, which cannot withstand significant salinization. This Special Issue looks at the roles that can be played by halophytes, extremophiles that do tolerate salinities toxic to most plants. In an ecological context, papers deal with the conservation of a rare species, the effects of rising concentrations of CO2 and flooding on coastal vegetation, and the consequences of tree planting in inland plains for salinization. Physiological studies deal with the different effects of chlorides and sulfates on the growth of halophytes, the ability of some parasitic plants to develop succulence when growing on halophytic hosts and the interesting finding that halophytes growing in their natural habitat do not show signs of oxidative stress. Nevertheless, spraying with ascorbic acid can enhance ascorbic acid-dependent antioxidant enzymes and growth in a species of Limonium. Enzymes preventing oxidative stress are expressed constitutively as is the case with the vacuolar H-ATPase, a key enzyme in ion compartmentation. A comparison of salt-excreting and non-excreting grasses showed the former to have higher shoot to root Na+ ratios than the latter. A particularly tolerant turf grass is described, as is the significance of its ability to secrete ions. A study of 38 species showed the importance of the interaction of a low osmotic potential and cell wall properties in maintaining growth. From an applied point of view, the importance of identifying genotypes and selecting those best suited for the product required, optimizing the conditions necessary for germination and maximizing yield are described. The consequence of selection for agronomic traits on salt tolerance is evaluated, as is the use of halophytes as green manures. Halophytes are remarkable plants: they are rare in relation to the total number of flowering plants and they tolerate salinities that most species cannot. It is clear from the papers published in this Special Issue that research into halophytes has a distinct place in aiding our understanding of salt tolerance in plants, an understanding that is likely to be of importance as climate change and population growth combine to challenge our ability to feed the human population of the world.
doi:10.1093/aobpla/plv020
PMCID: PMC4422832  PMID: 25757984
Climate change; halophyte; salinity; salt
8.  Influence of sulfur and cadmium on antioxidants, phytochelatins and growth in Indian mustard 
AoB Plants  2015;7:plv001.
Sulphur(S)-deficiency is emerging as a major problem for agricultural productivity. Cadmium (Cd) exerts its phytotoxicity against defence, growth and development. S-rich compounds (glutathione, phytochelatins, etc.) limit the impacts of Cd-toxicity. We investigated what happens during S-deficiency and Cd exposure (dual stress) in mustard. Major findings were: S-deficiency increases the susceptibility of plants to Cd-generated oxidative damage and modulates the AsA-GSH antioxidant cycle; SOD is not the first line of defence against metal stress and S-rich metabolites play a prime role; S-deprived plants are more prone to Cd and oxidative stress; and great loss is incurred to defence modules and growth under dual stress, restricting the efficiency of phytoremediation.
Soils in many parts of the world are contaminated with heavy metals, leading to multiple, deleterious effects on plants and threats to world food production efficiency. Cadmium (Cd) is one such metal, being toxic at relatively low concentrations as it is readily absorbed and translocated in plants. Sulfur-rich compounds are critical to the impact of Cd toxicity, enabling plants to increase their cellular defence and/or sequester Cd into vacuoles mediated by phytochelatins (PCs). The influence of sulfur on Cd-induced stress was studied in the hyperaccumulator plant Indian mustard (Brassica juncea) using two sulfur concentrations (+S, 300 µM SO42− and S-deficient −S, 30μMSO42−) with and without the addition of Cd (100 µM CdCl2) at two different time intervals (7 and 14 days after treatment). Compared with control plants (+S/−Cd), levels of oxidative stress were higher in S-deficient (−S/−Cd) plants, and greatest in S-deficient Cd-treated (−S/+Cd) plants. However, additional S (+S/+Cd) helped plants cope with oxidative stress. Superoxide dismutase emerged as a key player against Cd stress under both −S and +S conditions. The activity of ascorbate peroxidase, glutathione reductase and catalase declined in Cd-treated and S-deficient plants, but was up-regulated in the presence of sulfur. Sulfur deficiency mediated a decrease in ascorbate and glutathione (GSH) content but changes in ascorbate (reduced : oxidized) and GSH (reduced : oxidized) ratios were alleviated by sulfur. Our data clearly indicate that a sulfur pool is needed for synthesis of GSH, non-protein thiols and PCs and is also important for growth. Sulfur-based defence mechanisms and the cellular antioxidant pathway, which are critical for tolerance and growth, collapsed as a result of a decline in the sulfur pool.
doi:10.1093/aobpla/plv001
PMCID: PMC4323519  PMID: 25587194
Antioxidants; cadmium; growth; oxidative stress; phytochelatins; sulfur
9.  Effects of salinity and ascorbic acid on growth, water status and antioxidant system in a perennial halophyte 
AoB Plants  2015;7:plv004.
Limonium stocksii is a potential commercial cut-flower crop for saline areas using brackish water. We therefore were interested to learn about the mechanism of its salinity tolerance. Plants grew well under lower saline conditions (300 mM NaCl) but higher salinities reduced growth. An increase in leaf osmolality and the management of salinity-induced oxidative stress are the key strategies employed. Exogenous AsA application improved the functioning of the AsA-dependent antioxidant system, leading to better growth.
Salinity causes oxidative stress in plants by enhancing production of reactive oxygen species, so that an efficient antioxidant system, of which ascorbic acid (AsA) is a key component, is an essential requirement of tolerance. However, antioxidant responses of plants to salinity vary considerably among species. Limonium stocksii is a sub-tropical halophyte found in the coastal marshes from Gujarat (India) to Karachi (Pakistan) but little information exists on its salt resistance. In order to investigate the role of AsA in tolerance, 2-month-old plants were treated with 0 (control), 300 (moderate) and 600 (high) mM NaCl for 30 days with or without exogenous application of AsA (20 mM) or distilled water. Shoot growth of unsprayed plants at moderate salinity was similar to that of controls while at high salinity growth was inhibited substantially. Sap osmolality, AsA concentrations and activities of AsA-dependant antioxidant enzymes increased with increasing salinity. Water spray resulted in some improvement in growth, indicating that the growth promotion by exogenous treatments could partly be attributed to water. However, exogenous application of AsA on plants grown under saline conditions improved growth and AsA dependent antioxidant enzymes more than the water control treatment. Our data show that AsA-dependent antioxidant enzymes play an important role in salinity tolerance of L. stocksii.
doi:10.1093/aobpla/plv004
PMCID: PMC4334656  PMID: 25603966
Antioxidant; exogenous ascorbic acid application; halophyte; Limonium stocksii; salt tolerance
10.  Effects of disturbance on vegetation by sand accretion and erosion across coastal dune habitats on a barrier island 
AoB Plants  2015;7:plv003.
Understanding how plants stabilize coastal dunes will be important for predicting effects of climate change along shorelines. This study used three years of data on plant communities and dune morphology on a barrier island to show how particular plant communities were associated with the disturbances that occur in coastal dunes. As expected, foredunes were very disturbed, with important effects on their vegetation. However, the much smaller disturbances that occurred in areas behind the foredunes had similar effects on the plant communities. These results will be important for modeling how increased storms and ocean level rise will affect sandy coasts.
Coastal geomorphology and vegetation are expected to be particularly sensitive to climate change, because of disturbances caused by sea-level rise and increased storm frequency. Dunes have critical reciprocal interactions with vegetation; dunes create habitats for plants, while plants help to build dunes and promote geomorphological stability. These interactions are also greatly affected by disturbances associated with sand movement, either in accretion (dune building) or in erosion. The magnitude and intensity of disturbances are expected to vary with habitat, from the more exposed and less stable foredunes, to low-lying and flood-prone interdunes, to the protected and older backdunes. Permanent plots were established at three different spatial scales on St George Island, FL, USA, where the vegetation and dune elevation were quantified annually from 2011 to 2013. Change in elevation, either through accretion or erosion, was used as a measure of year-to-year disturbance over the 2 years of the study. At the scale of different dune habitats, foredunes were found to have the greatest disturbance, while interdunes had the least. Elevation and habitat (i.e. foredune, interdune, backdune) were significantly correlated with plant community composition. Generalized linear models conducted within each habitat show that the change in elevation (disturbance) is also significantly correlated with the plant community, but only within foredunes and interdunes. The importance of disturbance in exposed foredunes was expected and was found to be related to an increasing abundance of a dominant species (Uniola paniculata) in eroding areas. The significant effect of disturbance in the relatively stable interdunes was surprising, and may be due to the importance of flooding associated with small changes in elevation in these low-lying areas. Overall, this study documents changes in the plant community associated with elevation, and demonstrates that the foredune and interdune communities are also associated with the responses of specific species to local changes in elevation due to accretion or erosion.
doi:10.1093/aobpla/plv003
PMCID: PMC4340151  PMID: 25587195
Climate change; coastal zones; disturbance; geomorphology; ordination; plant community
11.  Suppression of annual Bromus tectorum by perennial Agropyron cristatum: roles of soil nitrogen availability and biological soil space 
AoB Plants  2015;7:plv006.
Sagebrush (Artemisia) rangelands of the western United States are becoming dominated by the exotic annual grass cheatgrass (Bromus tectorum). Rehabilitation of invaded rangelands is predicated on establishing healthy and dense perennial grass communities, which suppress cheatgrass. Our research investigated how established plants of the perennial grass, crested wheatgrass (Agropyron cristatum), suppress cheatgrass. Our data suggest that established crested wheatgrass reduces soil nitrogen availability and occupies biological soil space such that growth of cheatgrass is significantly reduced. Greater understanding of the role of biological soil space could be used to breed and select plant materials with traits that are more suppressive to invasive annual grasses.
Worldwide, exotic invasive grasses have caused numerous ecosystem perturbations. Rangelands of the western USA have experienced increases in the size and frequency of wildfires largely due to invasion by the annual grass Bromus tectorum. Rehabilitation of invaded rangelands is difficult; but long-term success is predicated on establishing healthy and dense perennial grass communities, which suppress B. tectorum. This paper reports on two experiments to increase our understanding of soil factors involved in suppression. Water was not limiting in this study. Growth of B. tectorum in soil conditioned by and competing with the exotic perennial Agropyron cristatum was far less relative to its growth without competition. When competing with A. cristatum, replacing a portion of conditioned soil with fresh soil before sowing of B. tectorum did not significantly increase its growth. The ability of conditioned soil to suppress B. tectorum was lost when it was separated from growing A. cristatum. Soil that suppressed B. tectorum growth was characterized by low mineral nitrogen (N) availability and a high molar ratio of NO2− in the solution-phase pool of NO2−+NO3−. Moreover, resin availability of NO2−+NO3− explained 66 % of the variability in B. tectorum above-ground mass, attesting to the importance of A. cristatum growth in reducing N availability to B. tectorum. Trials in which B. tectorum was suppressed the most were characterized by very high shoot/root mass ratios and roots that have less root hair growth relative to non-suppressed counterparts, suggesting co-opting of biological soil space by the perennial grass as another suppressive mechanism. Greater understanding of the role of biological soil space could be used to breed and select plant materials with traits that are more suppressive to invasive annual grasses.
doi:10.1093/aobpla/plv006
PMCID: PMC4340153  PMID: 25603967
Plant–soil relationships; root competition
12.  Long-distance plant dispersal to North Atlantic islands: colonization routes and founder effect 
AoB Plants  2015;7:plv036.
Our study provides new knowledge of two processes that are important for plant adaptation in a changing environment: 1) long-distance dispersal patterns, and 2) genetic founder effect on islands. Although the theoretical framework for the genetic founder effect on islands was proposed in 1973, we are the first to quantify it in relation to island size, dispersal distance, and plant traits. In addition, our genetic results are mainly coherent with post-glacial colonisation rather than in situ glacial survival, and should therefore bring a final end to the 140-year-long glacial survival-tabula rasa debate among northern biologists.
Long-distance dispersal (LDD) processes influence the founder effect on islands. We use genetic data for 25 Atlantic species and similarities among regional floras to analyse colonization, and test whether the genetic founder effect on five islands is associated with dispersal distance, island size and species traits. Most species colonized postglacially via multiple dispersal events from several source regions situated 280 to >3000 km away, and often not from the closest ones. A strong founder effect was observed for insect-pollinated mixed maters, and it increased with dispersal distance and decreased with island size in accordance with the theory of island biogeography. Only a minor founder effect was observed for wind-pollinated outcrossing species. Colonization patterns were largely congruent, indicating that despite the importance of stochasticity, LDD is mainly determined by common factors, probably dispersal vectors. Our findings caution against a priori assuming a single, close source region in biogeographic analyses.
doi:10.1093/aobpla/plv036
PMCID: PMC4432000  PMID: 25876627
Amplified fragment length polymorphism (AFLP); dispersal vector; founder effect; genetic diversity; islands; long-distance dispersal (LDD); postglacial; species traits
13.  Diversity of sexual systems within different lineages of the genus Silene 
AoB Plants  2015;7:plv037.
Species or populations may be categorized by sexual system. Here, we examine the frequency of sexual systems in Silene. We found that hermaphroditism is the most common sexual system, followed by dioecy, gynodioecy and gynodioecy-gynomonoecy (females, hermaphrodites and gynomonoecious plants) with similar frequency. These sexual systems are equally represented in the two phylogenetically supported subgenera Silene and Behenantha. We specifically studied the sexual systems of section Psammophilae (four species and 26 populations), and found that most populations are gynodioecious-gynomonoecious. Hermaphrodites are the most common sexual morph, and females tend to produce fewer flowers than other morphs.
Species and populations can be categorized by their sexual systems, depending on the spatial distribution of female and male reproductive structures within and among plants. Although a high diversity of sexual systems exists in Silene, their relative frequency at the genus and infrageneric level is unknown. Here, we carried out an extensive literature search for direct or indirect descriptions of sexual systems in Silene species. We found descriptions of sexual systems for 98 Silene species, where 63 and 35 correspond to the phylogenetically supported subgenera Silene and Behenantha, respectively. Hermaphroditism was the commonest sexual system (58.2 %), followed by dioecy (14.3 %), gynodioecy (13.3 %) and gynodioecy–gynomonoecy (i.e. hermaphroditic, female and gynomonoecious plants coexisting in the same population; 12.2 %). The presence of these sexual systems in both subgenera suggests their multiple origins. In 17 species, the description of sexual systems varied, and in most cases these differences corresponded to variations within or among populations. Interestingly, the poorly studied gynodioecy–gynomonoecy sexual system showed similar frequency to dioecy and gynodioecy in both subgenera. In addition, the incidence of gynodioecy–gynomonoecy was analysed in the species of section Psammophilae (Silene littorea, S. psammitis, S. adscendens and S. cambessedesii), in a survey of 26 populations across the distribution area of the species. The four species showed gynomonoecy–gynodioecy in most populations. Hermaphrodites were the most frequent morph, with a low number of females and gynomonoecious plants in all populations. The frequency of sexual morphs varied significantly among the studied populations but not among species. Female plants generally produced smaller numbers of flowers than hermaphroditic or gynomonoecious plants, and the percentages of female flowers per population were low. All these findings suggest that the gynodioecious–gynomonoecious sexual system in section Psammophilae is closer to hermaphroditism or gynomonoecy than gynodioecy.
doi:10.1093/aobpla/plv037
PMCID: PMC4433491  PMID: 25862920
Behenantha; Caryophyllaceae; dioecy; gynodioecy; gynodioecy–gynomonoecy; hermaphroditism; Psammophilae; sexual polymorphism; sexual system; Silene
14.  Genomic sequencing and microsatellite marker development for Boswellia papyrifera, an economically important but threatened tree native to dry tropical forests 
AoB Plants  2015;7:plu086.
The world famous aromatic resin Frankincense is tapped from natural populations of Boswellia trees. Most of these populations have been shrinking rapidly over recent decades. To help guide conservation efforts for imperilled species of this genus, we developed 46 genetic markers for Boswellia papyrifera. Several of these were cross-transferable to other Boswellia species that occur in Ethiopia and Yemen. We also identified genes involved in the biosynthesis of the terpenes and terpenoids that are major constituents of frankincense.
Microsatellite (or simple sequence repeat, SSR) markers are highly informative DNA markers often used in conservation genetic research. Next-generation sequencing enables efficient development of large numbers of SSR markers at lower costs. Boswellia papyrifera is an economically important tree species used for frankincense production, an aromatic resinous gum exudate from bark. It grows in dry tropical forests in Africa and is threatened by a lack of rejuvenation. To help guide conservation efforts for this endangered species, we conducted an analysis of its genomic DNA sequences using Illumina paired-end sequencing. The genome size was estimated at 705 Mb per haploid genome. The reads contained one microsatellite repeat per 5.7 kb. Based on a subset of these repeats, we developed 46 polymorphic SSR markers that amplified 2–12 alleles in 10 genotypes. This set included 30 trinucleotide repeat markers, four tetranucleotide repeat markers, six pentanucleotide markers and six hexanucleotide repeat markers. Several markers were cross-transferable to Boswellia pirrotae and B. popoviana. In addition, retrotransposons were identified, the reads were assembled and several contigs were identified with similarity to genes of the terpene and terpenoid backbone synthesis pathways, which form the major constituents of the bark resin.
doi:10.1093/aobpla/plu086
PMCID: PMC4433549  PMID: 25573702
Conservation genetics; resin; SSR; terpene biosynthesis; terpenoid; tropical dry forest.
15.  Growth and nitrogen fixation of legumes at increased salinity under field conditions: implications for the use of green manures in saline environments 
AoB Plants  2015;7:plv010.
In this paper we identify yellow sweet clover (Melilotus officinalis) as a suitable candidate to be adopted as a green manure in saline agriculture. The plants perform well up to a third seawater salinity concentration and continue to get their nitrogen from symbiotic root bacteria. Alternatively, alfalfa (Medicago sativa) could be used since it shows similar tolerance to salinity. However, since the absolute biomass of yellow sweet clover is considerably higher, this plant would be our choice if the goal is to enrich soils with organic material and, specifically, with a sustainable input of nitrogen.
The use of legumes as green manure can potentially increase crop productivity in saline environments and thus contribute to the sustainability of agricultural systems. Here, we present results from a field experiment conducted in the Netherlands that addressed the efficiency of nitrogen (N) fixation by a legume at varying salinities. We grew Melilotus officinalis in an agricultural field using drip irrigation with water salinity varying in electrical conductivity between 1.7 and 20 dS m−1. In the experiment, nearly 100 % of total plant N in M. officinalis was derived from symbiotic fixation at all but the highest salinity level (20 dS m−1). Our results indicated that this species derived substantial amounts of N via symbiotic fixation, the N becoming available in the soil (and thus available to crops) when cultivated legumes senesce and decompose. Based on the growth performance of M. officinalis and its ability to fix N at moderate soil salinity in our field experiments, we identified this species as a promising source for green manure in saline agriculture in temperate regions.
doi:10.1093/aobpla/plv010
PMCID: PMC4371099  PMID: 25661201
Halophytes; Melilotus officinalis; salinity; Sesbania; symbiotic nitrogen fixation
16.  Variation in reward quality and pollinator attraction: the consumer does not always get it right 
AoB Plants  2015;7:plv034.
Bees depend on pollen as the primary protein source for their larvae and should be strongly selected to identify cues associated with the most rewarding flowers. We examined the ability of bumble bees (Bombus impatiens) to identify the most rewarding foraging opportunities using arrays of live monkeyflowers (Mimulus guttatus), artificial plants, and pairwise olfactory tests. Bees could identify pollen rewards by scent and tended to visit the most rewarding artificial flowers. They seemed less able to identify the best pollen sources when foraging on live plants. We suggest that live plants may provide conflicting or deceptive signals to pollinators.
Nearly all bees rely on pollen as the sole protein source for the development of their larvae. The central importance of pollen for the bee life cycle should exert strong selection on their ability to locate the most rewarding sources of pollen. Despite this importance, very few studies have examined the influence of intraspecific variation in pollen rewards on the foraging decisions of bees. Previous studies have demonstrated that inbreeding reduces viability and hence protein content in Mimulus guttatus (seep monkeyflower) pollen and that bees strongly discriminate against inbred in favour of outbred plants. We examined whether variation in pollen viability could explain this preference using a series of choice tests with living plants, artificial plants and olfactometer tests using the bumble bee Bombus impatiens. We found that B. impatiens preferred to visit artificial plants provisioned with fertile anthers over those provisioned with sterile anthers. They also preferred fertile anthers when provided only olfactory cues. These bumble bees were unable to discriminate among live plants from subpopulations differing dramatically in pollen viability, however. They preferred outbred plants even when those plants were from subpopulations with pollen viability as low as the inbred populations. Their preference for outbred plants was evident even when only olfactory cues were available. Our data showed that bumble bees are able to differentiate between anthers that provide higher rewards when cues are isolated from the rest of the flower. When confronted with cues from the entire flower, their choices are independent of the quality of the pollen reward, suggesting that they are responding more strongly to cues unassociated with rewards than to those correlated with rewards. If so, this suggests that a sensory bias or some level of deception may be involved with advertisement to pollinators in M. guttatus.
doi:10.1093/aobpla/plv034
PMCID: PMC4417137  PMID: 25858692
Bombus impatiens; floral reward; honest signal; inbreeding; Mimulus guttatus; olfaction; pollen quality; pollinator preference
17.  Effects of a native parasitic plant on an exotic invader decrease with increasing host age 
AoB Plants  2015;7:plv031.
Infection by parasitic plants has been considered as an effective method for controlling invasive plants because the parasites partially or completely absorb water, nutrients, and carbohydrates from their host plants, suppressing the vitality of the host. Our study verified that younger and smaller Bidens pilosa plants suffer from higher levels of damage and are less likely to recover from infection by the parasitic plant Cuscuta australis than relatively older and larger plants, suggesting that Cuscuta australis is only a viable biocontrol agent for younger Bidens pilosa plants.
Understanding changes in the interactions between parasitic plants and their hosts in relation to ontogenetic changes in the hosts is crucial for successful use of parasitic plants as biological controls. We investigated growth, photosynthesis and chemical defences in different-aged Bidens pilosa plants in response to infection by Cuscuta australis. We were particularly interested in whether plant responses to parasite infection change with changes in the host plant age. Compared with the non-infected B. pilosa, parasite infection reduced total host biomass and net photosynthetic rates, but these deleterious effects decreased with increasing host age. Parasite infection reduced the concentrations of total phenolics, total flavonoids and saponins in the younger B. pilosa but not in the older B. pilosa. Compared with the relatively older and larger plants, younger and smaller plants suffered from more severe damage and are likely less to recover from the infection, suggesting that C. australis is only a viable biocontrol agent for younger B. pilosa plants.
doi:10.1093/aobpla/plv031
PMCID: PMC4417202  PMID: 25838325
Defence; deleterious effect; growth; invasive plant; parasitic plant
18.  Competition among native and invasive Impatiens species: the roles of environmental factors, population density and life stage 
AoB Plants  2015;7:plv033.
We examined competition effects in an experiment with three Impatiens species (Balsaminaceae) sharing similar life-history characteristics and habitats: the native I. noli-tangere, and two invasive species, I. parviflora and I. glandulifera. The results suggest that the effect of competition on the performance of invasive Impatiens species exceeds that of environmental factors, i.e. light and soil moisture. Competitive interactions with co-occurring congeners may thus be a more important predictor of the invasion success of an invasive species and its population dynamics than its response to abiotic factors, and should be taken into account when evaluating their invasion potential.
Many invasive species are considered competitively superior to native species, with the strongest competition expected in species with similar niches and/or in closely related species. However, competition outcome is strongly context-dependent as competitive strength varies along environmental gradients, and life stages, and also depends on abundances. To explore the importance of these factors, we examined competition effects in an experiment with three Impatiens species (Balsaminaceae) widespread in central Europe and sharing similar life-history characteristics and habitats: the native I. noli-tangere, and two invasive species, I. parviflora and I. glandulifera. We compared their competitive strength and reciprocal impacts under two levels of water and light availability, two overall planting densities and three competitor densities. We assessed species performance (ability to complete the life-cycle, biomass and fecundity) and temporal competition dynamics in a garden pot experiment. Environmental variables had lower explanatory power than overall planting and competitor density, which indicates the importance of competitive interactions when evaluating plant performance and potential invasion success. Despite poor and delayed germination, the invasive I. glandulifera attained dominance even at a high competitor density and was competitively superior across all treatments, exceeding the height of both congeners. Impatiens parviflora was competitively weakest, having a negligible impact on both native I. noli-tangere and invasive I. glandulifera. The intermediate competitive strength of the native I. noli-tangere probably results from its intermediate height, and good germination rate and timing. The difference in height among species increased during the experiment when I. glandulifera was involved; this species continues growing until autumn, enhancing its competitive superiority. The results provide a mechanistic understanding for the competitive exclusion of native I. noli-tangere that occurs in stands with I. glandulifera, but the limited impact of I. parviflora on I. noli-tangere in their mixed stands.
doi:10.1093/aobpla/plv033
PMCID: PMC4417208  PMID: 25832103
Alien species; balsam; competition; congeners; plant density; shading levels; water availability
19.  Dominant plant taxa predict plant productivity responses to CO2 enrichment across precipitation and soil gradients 
AoB Plants  2015;7:plv027.
Continued increases in atmospheric carbon dioxide will increase the productivity of rangelands partly by plant community change. Soil moisture increases may contribute more in arid systems or on coarser-textured soils, but not in deserts. Surprisingly, community change explained virtually all of yearly productivity increases in semi-arid grassland and up to 80% in mesic grassland, but only on some soil types, and a xeric shrubland exhibited no productivity, soil moisture or community change responses. Thus, community change dominated the productivity response to CO2 enrichment in grasslands, and the contribution of yearly soil moisture responses requires further research.
The Earth's atmosphere will continue to be enriched with carbon dioxide (CO2) over the coming century. Carbon dioxide enrichment often reduces leaf transpiration, which in water-limited ecosystems may increase soil water content, change species abundances and increase the productivity of plant communities. The effect of increased soil water on community productivity and community change may be greater in ecosystems with lower precipitation, or on coarser-textured soils, but responses are likely absent in deserts. We tested correlations among yearly increases in soil water content, community change and community plant productivity responses to CO2 enrichment in experiments in a mesic grassland with fine- to coarse-textured soils, a semi-arid grassland and a xeric shrubland. We found no correlation between CO2-caused changes in soil water content and changes in biomass of dominant plant taxa or total community aboveground biomass in either grassland type or on any soil in the mesic grassland (P > 0.60). Instead, increases in dominant taxa biomass explained up to 85 % of the increases in total community biomass under CO2 enrichment. The effect of community change on community productivity was stronger in the semi-arid grassland than in the mesic grassland, where community biomass change on one soil was not correlated with the change in either the soil water content or the dominant taxa. No sustained increases in soil water content or community productivity and no change in dominant plant taxa occurred in the xeric shrubland. Thus, community change was a crucial driver of community productivity responses to CO2 enrichment in the grasslands, but effects of soil water change on productivity were not evident in yearly responses to CO2 enrichment. Future research is necessary to isolate and clarify the mechanisms controlling the temporal and spatial variations in the linkages among soil water, community change and plant productivity responses to CO2 enrichment.
doi:10.1093/aobpla/plv027
PMCID: PMC4429605  PMID: 25829380
Central Plains grasslands; climate change; community change; Mojave Desert; primary productivity; rangelands; threshold responses
20.  Investigating the impacts of recycled water on long-lived conifers 
AoB Plants  2015;7:plv035.
This research has direct implications for public and private institutions seeking to conserve water by irrigating landscapes with recycled (a.k.a. reclaimed) water. Although typical salt contents in recycled water are low (< 2.0 dS m−1), levels may still be harmful to salt-sensitive plants. We discovered that salt accumulation in soils would negatively impact coast redwoods when recycled water salinity exceeds>1.0 dS m−1. This is the first paper reporting the impacts of salinity on the growth of the coast redwood. The results suggest that irrigation management of long-lived conifers will be essential to protect these important trees.
Recycled wastewater is a popular alternative water resource. Recycled water typically has higher salinity than potable water and therefore may not be an appropriate water source for landscapes planted with salt-intolerant plant species. Coast redwoods (Sequoia sempervirens) are an important agricultural, horticultural and ecological species assumed to be salt intolerant. However, no studies have analysed how salinity impacts coast redwood growth. To determine salt-related growth limitations, as well as susceptibility to particular salt ions, we divided 102 S. sempervirens ‘Aptos Blue’ saplings evenly into 17 salinity treatments: a control and four different salts (sodium chloride, calcium chloride, sodium chloride combined with calcium chloride, and sodium sulfate). Each salt type was applied at four different concentrations: 1.0, 3.0, 4.5 and 6.0 dS m−1. Trees were measured for relative growth, and leaves were analysed for ion accumulation. Results showed that the relative stem diameter growth was inversely proportional to the increase in salinity (electrical conductivity), with R2 values ranging from 0.72 to 0.82 for different salts. Analysis of variance tests indicated that no particular salt ion significantly affected growth differently than the others (P > 0.1). Pairwise comparisons of the means revealed that moderately saline soils (4–8 dS m−1) would decrease the relative height growth by 30–40 %. Leaf tissue analysis showed that all treatment groups accumulated salt ions. This finding suggests reduced growth and leaf burn even at the lowest ion concentrations if salts are not periodically leached from the soil. Regardless of the specific ions in the irrigation water, the results suggest that growth and appearance of coast redwoods will be negatively impacted when recycled water electrical conductivity exceeds >1.0 dS m−1. This information will prove valuable to many metropolitan areas faced with conserving water while at the same time maintaining healthy verdant landscapes that include coast redwoods and other long-lived conifers.
doi:10.1093/aobpla/plv035
PMCID: PMC4455786  PMID: 25876628
California; drought; Mediterranean climate; reclaimed water; urban forestry; urban horticulture
21.  Tolerance of three grain legume species to transient waterlogging 
AoB Plants  2015;7:plv040.
Crop production can be limited by soil waterlogging. Tolerance to waterlogging can vary between and within species. This study quantified tolerance to soil waterlogging in two divergent genotypes of pea (Pisum sativum), two of lentil (Lens culinaris) and a grasspea (Lathyrus sativus) control at germination and during vegetative growth. Soil waterlogging at 10 mm depth had no significant effect on shoot and root dry mass after 14 days. Significant genetic variation in both pea and lentil in tolerance to waterlogging after germination and subsequent recovery was evident. Screening of additional pea and lentil germplasm for waterlogging conditions is clearly warranted.
Waterlogging reduces the yield of food crops. Tolerance to waterlogging could vary between and within species. This study aimed to quantify tolerance to soil waterlogging in two divergent genotypes of pea (Pisum sativum), two of lentil (Lens culinaris) and a grasspea (Lathyrus sativus) control at germination and during vegetative growth. Following germination, seeds were grown for 14 days in soil waterlogged with the water table 10 mm below the surface, and then by draining the pots and allowing to recover for 21 days—to be compared with 35 days of continuous waterlogging. In both pea and lentil, the pair of genotypes contrasted widely with large-seeded pea genotype Kaspa and lentil genotype Nugget showing higher (2-fold) root porosity and less effect on shoot nitrogen content under waterlogging than the other genotypes (NPE and ATC). During recovery, the same two genotypes—Kaspa pea and Nugget lentil—also recovered better than their smaller-seeded species pairs. Soil waterlogging at 10 mm depth had no significant effect on shoot and root dry mass after 14 days. Root penetration into waterlogged soil was restricted to ∼100 mm depth and its distribution altered for pea and lentil genotypes but not for grasspea. Within the small sample studied, we demonstrated a significant genetic variation in both pea and lentil in tolerance to waterlogging after germination and subsequent recovery for the first time. Screening of additional pea and lentil germplasm for waterlogging conditions is clearly warranted.
doi:10.1093/aobpla/plv040
PMCID: PMC4455828  PMID: 25902834
Chlorophyll; intra-species variation; legume; porosity; waterlogging
22.  Extractable nitrogen and microbial community structure respond to grassland restoration regardless of historical context and soil composition 
AoB Plants  2015;7:plu085.
Exotic plants have the capacity to establish positive plant-soil feedbacks facilitating invader persistence often at the cost of native plant species success. We compared the plant communities, soil chemical and microbial communities and nutrient turnover rates between invaded and restored plots in inland and coastal grasslands. Greater extractable N, slower N cycling rates and differing microbial community composition resulted from restoration. These differences indicate that grassland soils are responsive to vegetation change, and therefore not resistant to invasion-caused feedbacks. It also suggests these soils are resilient to invasion and may be capable of recovery following vegetative restoration.
Grasslands have a long history of invasion by exotic annuals, which may alter microbial communities and nutrient cycling through changes in litter quality and biomass turnover rates. We compared plant community composition, soil chemical and microbial community composition, potential soil respiration and nitrogen (N) turnover rates between invaded and restored plots in inland and coastal grasslands. Restoration increased microbial biomass and fungal : bacterial (F : B) ratios, but sampling season had a greater influence on the F : B ratio than did restoration. Microbial community composition assessed by phospholipid fatty acid was altered by restoration, but also varied by season and by site. Total soil carbon (C) and N and potential soil respiration did not differ between treatments, but N mineralization decreased while extractable nitrate and nitrification and N immobilization rate increased in restored compared with unrestored sites. The differences in soil chemistry and microbial community composition between unrestored and restored sites indicate that these soils are responsive, and therefore not resistant to feedbacks caused by changes in vegetation type. The resilience, or recovery, of these soils is difficult to assess in the absence of uninvaded control grasslands. However, the rapid changes in microbial and N cycling characteristics following removal of invasives in both grassland sites suggest that the soils are resilient to invasion. The lack of change in total C and N pools may provide a buffer that promotes resilience of labile pools and microbial community structure.
doi:10.1093/aobpla/plu085
PMCID: PMC4323520  PMID: 25555522
Carbon; exotic grasses; exotic plants; phospholipid fatty acid; resilience.
23.  The behavioural ecology of climbing plants 
AoB Plants  2015;7:plv013.
Climbing plants require an external support to grow vertically and thus achieve better access to sunlight. Climbing plants that find a suitable support have greater performance, size and reproduction than those that remain prostrate. Plant behaviour involves rapid morphological or physiological responses to events or environmental changes. Theoretical frameworks from behavioural ecology, traditionally applied to animals, have been successfully used to study plant behaviour. I herein review studies addressing ecological causes and consequences of support finding and use by climbing plants. I also propose the use of behavioural ecology theoretical frameworks to study climbing plant behaviour.
Climbing plants require an external support to grow vertically and enhance light acquisition. Vines that find a suitable support have greater performance and fitness than those that remain prostrate. Therefore, the location of a suitable support is a key process in the life history of climbing plants. Numerous studies on climbing plant behaviour have elucidated mechanistic details of support searching and attachment. Far fewer studies have addressed the ecological significance of support-finding behaviour and the factors that affect it. Without this knowledge, little progress can be made in the understanding of the evolution of support-finding behaviour in climbers. Here I review studies addressing ecological causes and consequences of support finding and use by climbing plants. I also propose the use of behavioural ecology theoretical frameworks to study climbing plant behaviour. I show how host tree attributes may determine the probability of successful colonization for the different types of climbers, and examine the evidence of environmental and genetic control of circumnutation behaviour and phenotypic responses to support availability. Cases of oriented vine growth towards supports are highlighted. I discuss functional responses of vines to the interplay between herbivory and support availability under different abiotic environments, illustrating with one study case how results comply with a theoretical framework of behavioural ecology originally conceived for animals. I conclude stressing that climbing plants are suitable study subjects for the application of behavioural–ecological theory. Further research under this framework should aim at characterizing the different stages of the support-finding process in terms of their fit with the different climbing modes and environmental settings. In particular, cost–benefit analysis of climbing plant behaviour should be helpful to infer the selective pressures that have operated to shape current climber ecological communities.
doi:10.1093/aobpla/plv013
PMCID: PMC4363473  PMID: 25678588
Behavioural ecology; circumnutation; climbing plants; lianas; optimal foraging; support-searching; vines
24.  Developmental stages of cultivated strawberry flowers in relation to chilling sensitivity 
AoB Plants  2015;7:plv012.
During strawberry flower development, a down-shift of temperatures below 2°C translates into injury to reproductive organs from early stages of bud development until the flower opens. Pollen development is especially vulnerable to chilling but carpels were also affected at the end of their maturation. By associating flower developmental stages to specific bud sizes, it was possible to identify when critical processes are taking place during flower development. These results can be used to shed light on susceptibility to low temperatures of other fruiting crops belonging to the Rosaceae.
Environmental factors affecting flower development may limit the yields of fruiting crops worldwide. In temperate regions, chilling temperatures during flower development can compromise fruit production, but their negative effects vary depending on the differing susceptibilities of each developmental stage. The cultivated strawberry (Fragaria× ananassa Duch.) is widely grown worldwide but financial returns are influenced by sudden shifts to chilling temperatures occurring during the cropping cycle. Despite this important limitation, knowledge of F.× ananassa flower development is lacking, in contrast to the diploid wild-type strawberry (F. vesca). In this study we describe steps in floral development of cultivated strawberry and define their vulnerability to chilling temperatures. To achieve this, flower buds from strawberry plants of cv. ‘Camarosa’ were labelled and monitored from bud initiation until anthesis. Description of morphological and functional changes during flower development was based on histological sections and scanning electron microscopy. To determine the impact of low temperatures at different developmental stages, plants carrying buds of different sizes were chilled at 2 °C for 24 h. Several parameters related to male and female gametophyte development were later evaluated in flowers as they approached anthesis. Fragaria× ananassa flower development was divided into 16 stages according to landmark events. These stages were similar to those documented for F. vesca but three new additional intermediate stages were described. Timing of developmental processes was achieved by correlating developmental staging with specific bud sizes and days before anthesis. Time to reach anthesis from early bud stages was 17–18 days. During this period, we detected four critical periods vulnerable to low temperatures. These were mostly related to male gametophyte development but also to injury to female organs at late developmental stages. These results provide the essential groundwork on floral biology of cultivated strawberry that is a prerequisite for successful comparative studies of cold tolerance among genotypes during flower formation.
doi:10.1093/aobpla/plv012
PMCID: PMC4363474  PMID: 25661200
Critical periods; flower development; flower differentiation; Fragaria × ananassa; low temperature; pistil; pollen
25.  Rangewide ploidy variation and evolution in Acacia senegal: a north–south divide? 
AoB Plants  2015;7:plv011.
A recent study of Acacia senegal showed a geographic pattern of genetic variation, which differentiated East and Southern African populations from those in the Sudano-Sahelian region. We build on this previous research to explore variation in DNA content using the flow cytometry method and chromosome number. A geographic north-south DNA content pattern was detected, reflecting the previous results. These results suggest that DNA content may also be important in elucidating the evolutionary history and distribution of the species. Use of external tissues of dried twigs in flow cytometry is new, and presents the opportunity to study numerous other dryland woody species.
Knowledge of rangewide variation in DNA content and ploidy level may be valuable in understanding the evolutionary history of a species. Recent studies of Acacia senegal report diploids and occasional tetraploids in the Sudano-Sahelian region of sub-Saharan Africa, but nothing is known about the overall extent of DNA ploidy variation within the species. In this study, we determine the DNA content and ploidy level of A. senegal across its native range, and explore whether the variation is related to its evolutionary and colonization history. We used propidium iodide flow cytometry (FCM) to estimate DNA content (2C value) and infer ploidy in 157 individuals from 54 populations on various tissues, using seeds, fresh leaves, dried leaves and twigs and herbarium specimens. The mean 2C DNA (pg ± s.d.) contents detected were 1.47 ± 0.09, 2.12 ± 0.02, 2.89 ± 0.12, and a single individual with 4.51 pg, corresponding to a polyploid series of diploid, triploid, tetraploid and hexaploid individuals. Diploids were confirmed by chromosome counts (2n = 2x = 26). Most populations (90.7 %) were of single ploidy level, while mixed ploidy populations (9.3 %) comprising mostly diploids (2x+3x, 2x+4x and 2x+6x) were restricted to the Sudano-Sahelian and Indian subcontinent regions, its northern range. The species is predominantly diploid, and no mixed ploidy populations were detected in east and southern Africa, its southern range. The geographic pattern of ploidy variation in conjunction with existing phylogeographic and phylogenetic data of the species suggests that polyploids have occurred multiple times in its evolutionary and recent colonization history, including contemporary ecological timescales. The successful use of external tissues of dried twigs in FCM is new, and presents the opportunity to study numerous other dryland woody species.
doi:10.1093/aobpla/plv011
PMCID: PMC4363475  PMID: 25680798
Acacia senegal; African acacias; diploids; DNA ploidy level; dried leaves and twigs; flow cytometry; hexaploids; polyploidy; tetraploids; triploids

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