Leaves may function as temperature sensors in the heterophylly of Rorippa aquatica (Brassicaceae)

Many plants show heterophylly, which is variation in leaf form within a plant owing to environmental change. The molecular mechanisms underlying heterophylly have recently been investigated in several plant species. However, little is known about how plants exhibiting heterophylly sense environmental cues. Here, we used Rorippa aquatica (Brassicaceae), which shows heterophylly, to investigate whether a single leaf can sense and transit changes in ambient temperature. The morphology of newly developed leaves after single-leaf warming treatment was significantly different from that of mock-treated control leaves, suggesting that leaves are sensing organs that mediate the responses to changes in ambient temperature in R. aquatica.     

Mycorrhizal colonisation in plants from intermittent aquatic habitats

Several plant species with amphibious characteristics from intermittent aquatic habitats were examined for colonisation with arbuscular mycorrhizal fungi (AM), dark septate endophytes (DSE) and the ratio of aerenchyma in root tissue. We studied submerged specimens of Alisma plantago-aquatica, Mentha aquatica, Myosotis scorpioides, Oenanthe fistulosa, Gratiola officinalis, Glyceria fluitans, Sium latifolium and Teucrium scordium. In the first four, we also examined the emerged growth forms, which were grown under experimental conditions. Roots of all species were mycorrhizal and showed AM and DSE colonisation. The results suggest that AM colonisation may also be abundant in plants of aquatic environment. Arbuscules were not found in submerged specimens of M. aquatica, O. fistulosa and S. latifolium. The AM colonisation was generally higher in emerged specimens as compared to submerged ones. The aerenchyma ratio in root cross-sections ranged from 10 to 50% and in most cases did not differ between submerged and emerged specimens. No clear relationship between AM colonisation and aerenchyma ratio was recognized, while a positive correlation between AM and plant available phosphorous was established.     

A Developmental Model for Branching Morphogenesis of Lake Cress Compound Leaf

Lake cress, Rorippa aquatica (Brassicaceae), is a semi-aquatic plant that exhibits a variety of leaf shapes, from simple leaves to highly branched compound leaves, depending on the environment. Leaf shape can vary within a single plant, suggesting that the variation can be explained by a simple model. In order to simulate the branched structure in the compound leaves of R. aquatica, we implemented reaction-diffusion (RD) patterning onto a theoretical framework that had been developed for serration distribution in the leaves of Arabidopsis thaliana, with the modification of the one-dimensional reaction-diffusion domain being deformed with the spatial periodicity of the RD pattern while expanding. This simple method using an iterative pattern could create regular and nested branching patterns. Subsequently, we verified the plausibility of our theoretical model by comparing it with the experimentally observed branching patterns. The results suggested that our model successfully predicted both the qualitative and quantitative aspects of the timing and positioning of branching in growing R. aquatica leaves.     

Differential genome evolution and speciation of Coix lacryma-jobi L. and Coix aquatica Roxb. hybrid guangxi revealed by repetitive sequence analysis and fine karyotyping

Abstract

Background

Coix, Sorghum and Zea are closely related plant genera in the subtribe Maydeae. Coix comprises 9–11 species with different ploidy levels (2n = 10, 20, 30, and 40). The exclusively cultivated C. lacryma-jobi L. (2n = 20) is widely used in East and Southeast Asia for food and medicinal applications. Three fertile cytotypes (2n = 10, 20, and 40) have been reported for C. aquatica Roxb. One sterile cytotype (2n = 30) closely related to C. aquatica has been recently found in Guangxi of China. This putative hybrid has been named C. aquatica HG (Hybrid Guangxi). The genome composition and the evolutionary history of C. lacryma-jobi and C. aquatica HG are largely unclear.

Results

About 76% of the genome of C. lacryma-jobi and 73% of the genome of C. aquatica HG are repetitive DNA sequences as shown by low coverage genome sequencing followed by similarity-based cluster analysis. In addition, long terminal repeat (LTR) retrotransposable elements are dominant repetitive sequences in these two genomes, and the proportions of many repetitive sequences in whole genome varied greatly between the two species, indicating evolutionary divergence of them. We also found that a novel 102 bp variant of centromeric satellite repeat CentX and two other satellites only appeared in C. aquatica HG. The results from FISH analysis with repeat probe cocktails and the data from chromosomes pairing in meiosis metaphase showed that C. lacryma-jobi is likely a diploidized paleotetraploid species and C. aquatica HG is possibly a recently formed hybrid. Furthermore, C. lacryma-jobi and C. aquatica HG shared more co-existing repeat families and higher sequence similarity with Sorghum than with Zea.

Conclusions

The composition and abundance of repetitive sequences are divergent between the genomes of C. lacryma-jobi and C. aquatica HG. The results from fine karyotyping analysis and chromosome pairing suggested diploidization of C. lacryma-jobi during evolution and C. aquatica HG is a recently formed hybrid. The genome-wide comparison of repetitive sequences indicated that the repeats in Coix were more similar to those in Sorghum than to those in Zea, which is consistent with the phylogenetic relationship reported by previous work.

Electronic supplementary material

The online version of this article (doi:10.1186/1471-2164-15-1025) contains supplementary material, which is available to authorized users.     

A Decrease in Ambient Temperature Induces Post-Mitotic Enlargement of Palisade Cells in North American Lake Cress

In order to maintain organs and structures at their appropriate sizes, multicellular organisms orchestrate cell proliferation and post-mitotic cell expansion during morphogenesis. Recent studies using Arabidopsis leaves have shown that compensation, which is defined as post-mitotic cell expansion induced by a decrease in the number of cells during lateral organ development, is one example of such orchestration. Some of the basic molecular mechanisms underlying compensation have been revealed by genetic and chimeric analyses. However, to date, compensation had been observed only in mutants, transgenics, and γ-ray–treated plants, and it was unclear whether it occurs in plants under natural conditions. Here, we illustrate that a shift in ambient temperature could induce compensation in Rorippa aquatica (Brassicaceae), a semi-aquatic plant found in North America. The results suggest that compensation is a universal phenomenon among angiosperms and that the mechanism underlying compensation is shared, in part, between Arabidopsis and R. aquatica.     

Regulation of the KNOX-GA Gene Module Induces Heterophyllic Alteration in North American Lake Cress

Plants show leaf form alteration in response to changes in the surrounding environment, and this phenomenon is called heterophylly. Although heterophylly is seen across plant species, the regulatory mechanisms involved are largely unknown. Here, we investigated the mechanism underlying heterophylly in Rorippa aquatica (Brassicaceae), also known as North American lake cress. R. aquatica develops pinnately dissected leaves in submerged conditions, whereas it forms simple leaves with serrated margins in terrestrial conditions. We found that the expression levels of KNOTTED1-LIKE HOMEOBOX (KNOX1) orthologs changed in response to changes in the surrounding environment (e.g., change of ambient temperature; below or above water) and that the accumulation of gibberellin (GA), which is thought to be regulated by KNOX1 genes, also changed in the leaf primordia. We further demonstrated that exogenous GA affects the complexity of leaf form in this species. Moreover, RNA-seq revealed a relationship between light intensity and leaf form. These results suggest that regulation of GA level via KNOX1 genes is involved in regulating heterophylly in R. aquatica. The mechanism responsible for morphological diversification of leaf form among species may also govern the variation of leaf form within a species in response to environmental changes.     

Germination ecology of two closely related taxa in the genus Oenanthe: Fine tuning for the habitat?

For two closely related amphibious plant taxa of the genus Oenanthe (Apiaceae) germination traits were examined. Habitats of the two taxa differ in hydroregime. The endemic Oenanthe conioides of the river Elbe estuary experiences daily tidal fluctuations whereas the widespread Oenanthe aquatica grows at the edge of ponds and in freshwater wetlands with rare and unpredictable fluctuations of water table.     

Characterization of the Cystoid Nematode Meloidoderita kirjanovae (Nemata: Sphaeronematidae) from Southern Italy

A population of the cystoid nematode Meloidoderita kirjanovae was detected parasitizing water mint (Mentha aquatica) in southern Italy. The morphological identification of this species was confirmed by molecular analysis using the internal transcribed spacer 1 (ITS1) and 5.8S gene sequences of nuclear ribosomal DNA (rDNA), which clearly separated it from the closely related species Meloidoderita polygoni. A phylogenetic analysis of M. kirjanovae with species of related genera was conducted using sequences of the D2-D3 expansion segments of the 28S nuclear ribosomal RNA gene. The resulting phylogenetic tree was congruent with trees from an extended dataset for Criconematina and Tylenchida. The basal position of the genus Meloidoderita together with Sphaeronema within the Criconematina clade in this tree may indicate their close relationships. The anatomical changes induced by M. kirjanovae population from Italy in water mint were similar to those reported for a nematode population infecting roots of M. longifolia in Israel. Nematode feeding caused the formation of a stellar syncytium that disorganized the pericycle and vascular root tissues.     

A hominoid distal tibia from the Miocene of Pakistan

A distal tibia, YGSP 1656, from the early Late Miocene portion of the Chinji Formation in Pakistan is described. The fossil is 11.4 million years old and is one of only six postcranial elements now assigned to Sivapithecus indicus. Aspects of the articular surface are cercopithecoid-like, suggesting some pronograde locomotor activities. However, YGSP 1656 possesses an anteroposteriorly compressed metaphysis and a mediolaterally thick medial malleolus, ape-like features functionally related to orthograde body postures and vertical climbing. YGSP 1656 lacks specializations found in the ankle of terrestrial cercopithecoids and thus Sivapithecus may have been primarily arboreal. Nevertheless, the morphology of this tibia is unique, consistent with other interpretations of Sivapithecus postcranial functional morphology that suggest the locomotion of this ape lacks a modern analog. Based on the limited postcranial remains from S. indicus, we hypothesize that this taxon exhibited substantial body size dimorphism.     

核盘菌侵染对拟南芥表皮蜡质结构及化学组成的影响

以野生型拟南芥与蜡质突变体cer1、cer4为试验材料,通过研究核盘菌胁迫对拟南芥茎表皮蜡质结构及组分含量的影响,揭示核盘菌侵染与表皮蜡质的关系。扫描电镜结果显示,野生型拟南芥蜡质晶体以垂直于表面的杆状、块状结构为主;突变体cer1晶体类型以水平的松针状、块状结构为主;突变体cer4蜡质晶体以垂直片层结构为主。核盘菌胁迫下,拟南芥蜡质晶体结构及分布形态发生变化。蜡质层结构在核盘菌胁迫下表现为:杆状、松针状蜡质晶体减少—蜡质晶体熔融—表皮"囊状凸起"—表皮膜层破裂。这些结构变化有利于病菌突破角质层屏障而侵入到植株体内。色质谱分析结果显示:与野生型相比,cer1突变体烷、次级醇、酮类显著减少;cer4突变体表现为一级醇含量减少。接种核盘菌后,野生型拟南芥与蜡质突变体一级醇类显著增加(cer1增加不显著);烷类、次级醇类、酮类含量与蜡质总量均显著减少,表明蜡质前体物质在受到核盘菌胁迫后更多地通过酰基还原途径生成一级醇,从而减少了由脱羰基途径所生成的蜡质组分。核盘菌通过改变表皮蜡质晶体结构与化学组分分泌量来促进侵染。     

Clonal plasticity of aquatic plant species submitted to mechanical stress: escape versus resistance strategy

Background and Aims

The plastic alterations of clonal architecture are likely to have functional consequences, as they affect the spatial distribution of ramets over patchy environments. However, little is known about the effect of mechanical stresses on the clonal growth. The aim of the present study was to investigate the clonal plasticity induced by mechanical stress consisting of continuous water current encountered by aquatic plants. More particularly, the aim was to test the capacity of the plants to escape this stress through clonal plastic responses.

Methods

The transplantation of ramets of the same clone in two contrasting flow velocity conditions was carried out for two species (Potamogeton coloratus and Mentha aquatica) which have contrasting clonal growth forms. Relative allocation to clonal growth, to creeping stems in the clonal biomass, number and total length of creeping stems, spacer length and main creeping stem direction were measured.

Key Results

For P. coloratus, plants exposed to water current displayed increased total length of creeping stems, increased relative allocation to creeping stems within the clonal dry mass and increased spacer length. For M. aquatica, plants exposed to current displayed increased number and total length of creeping stems. Exposure to current induced for both species a significant increase of the proportion of creeping stems in the downstream direction to the detriment of creeping stems perpendicular to flow.

Conclusions

This study demonstrates that mechanical stress from current flow induced plastic variation in clonal traits for both species. The responses of P. coloratus could lead to an escape strategy, with low benefits with respect to sheltering and anchorage. The responses of M. aquatica that may result in a denser canopy and enhancement of anchorage efficiency could lead to a resistance strategy.Key words: Phenotypic plasticity, morphology, submerged aquatic vegetation, clonality, clonal architecture, Potamogeton coloratus, Mentha aquatica, escape, resistance, mechanical stress, thigmomorphogenesis     

风毛菊属3种植物叶的解剖结构比较

采用石蜡切片法对分布于祁连山海拔5 000 m左右流石滩上菊科风毛菊属水母雪兔子（Saussurea medusa Maxim）、鼠曲雪兔子（Saussurea gnaphalodes (Royle) Sch.）、红叶雪兔子（Saussurea paxiana Diels.）3种植物叶片的解剖结构进行了比较研究,结果表明：叶片表皮细胞均为单层,上下表皮都有气孔分布,气孔不下陷;角质层较厚,叶表面均被单列细胞的表皮毛。3种植物均为异面叶;叶肉栅栏组织较发达,通常由2～3层细胞组成,但栅栏组织细胞排列较疏松;海绵组织存在大量的细胞间隙;叶肉中通气组织发达,且均有不规则裂生性气腔。叶脉维管束中韧皮部都具有异细胞存在。这些共同特征是3种植物对高山地区缺氧、低温、强辐射等自然条件长期适应的结果。但是,3种植物在叶片的外部形态特征、叶肉栅栏组织细胞的特点、维管束发育程度、内分泌结构、不规则裂生气腔等方面又存在明显的不同,表现出3种植物对环境的适应也是存在差异的。     

Leaf dimorphism in Thuja plicata and Platycladus orientalis (thujoid Cupressaceae s. str., Coniferales): the changes in morphology and anatomy from juvenile needle leaves to mature scale leaves

Thuja plicata and Platycladus orientalis initially produce only bifacial needle leaves. When the first lateral shoots develop, the leaf morphology and anatomy changes dramatically. Subsequently, only greatly reduced, bifacial scale leaves are developed. A new kind of “superimposed bifaciality” occurs with the change from juvenile needle leaves to mature scale leaves. Anatomical dorsiventrality affects not only the individual leaf, but also the complete plagiotropic lateral shoots of Thuja, which have a sun- and shade-exposed side. The upper light-exposed median leaves show adaxial leaf anatomy, contrary to the lower shaded median leaves showing abaxial leaf anatomy. Due to their mixed exposure, the lateral leaves show a lateral differentiation. At vertical lateral shoots of Platycladus, a predominant light-exposed side is absent. Thus, the anatomical dorsiventrality does not affect the complete shoot. Here the morphological abaxial side of a scale leaf becomes functionally and physiologically adaxial by reorientation of the palisade parenchyma and stomata. In juvenile needle leaves, the palisade parenchyma is located adaxial, with the majority of stomata being located abaxial. Conversely, in mature scale leaves, the palisade parenchyma is abaxial and the majority of stomata are adaxial.     

Development of Full-Length cDNAs from Chinese Cabbage (Brassica rapa Subsp. pekinensis) and Identification of Marker Genes for Defence Response

Arabidopsis belongs to the Brassicaceae family and plays an important role as a model plant for which researchers have developed fine-tuned genome resources. Genome sequencing projects have been initiated for other members of the Brassicaceae family. Among these projects, research on Chinese cabbage (Brassica rapa subsp. pekinensis) started early because of strong interest in this species. Here, we report the development of a library of Chinese cabbage full-length cDNA clones, the RIKEN BRC B. rapa full-length cDNA (BBRAF) resource, to accelerate research on Brassica species. We sequenced 10 000 BBRAF clones and confirmed 5476 independent clones. Most of these cDNAs showed high homology to Arabidopsis genes, but we also obtained more than 200 cDNA clones that lacked any sequence homology to Arabidopsis genes. We also successfully identified several possible candidate marker genes for plant defence responses from our analysis of the expression of the Brassica counterparts of Arabidopsis marker genes in response to salicylic acid and jasmonic acid. We compared gene expression of these markers in several Chinese cabbage cultivars. Our BBRAF cDNA resource will be publicly available from the RIKEN Bioresource Center and will help researchers to transfer Arabidopsis-related knowledge to Brassica crops.     

A new hero emerges: another exceptional mammalian spine and its potential adaptive significance

The hero shrew''s (Scutisorex somereni) massive interlocking lumbar vertebrae represent the most extreme modification of the vertebral column known in mammals. No intermediate form of this remarkable morphology is known, nor is there any convincing theory to explain its functional significance. We document a new species in the heretofore monotypic genus Scutisorex; the new species possesses cranial and vertebral features representing intermediate character states between S. somereni and other shrews. Phylogenetic analyses of DNA sequences support a sister relationship between the new species and S. somereni. While the function of the unusual spine in Scutisorex is unknown, it gives these small animals incredible vertebral strength. Based on field observations, we hypothesize that the unique vertebral column is an adaptation allowing these shrews to lever heavy or compressive objects to access concentrated food resources inaccessible to other animals.     

Feeding biomechanics in Acanthostega and across the fish–tetrapod transition

Acanthostega is one of the earliest and most primitive limbed vertebrates. Its numerous fish-like features indicate a primarily aquatic lifestyle, yet cranial suture morphology suggests that its skull is more similar to those of terrestrial taxa. Here, we apply geometric morphometrics and two-dimensional finite-element analysis to the lower jaws of Acanthostega and 22 other tetrapodomorph taxa in order to quantify morphological and functional changes across the fish–tetrapod transition. The jaw of Acanthostega is similar to that of certain tetrapodomorph fish and transitional Devonian taxa both morphologically (as indicated by its proximity to those taxa in morphospace) and functionally (as indicated by the distribution of stress values and relative magnitude of bite force). Our results suggest a slow tempo of morphological and biomechanical changes in the transition from Devonian tetrapod jaws to aquatic/semi-aquatic Carboniferous tetrapod jaws. We conclude that Acanthostega retained a primitively aquatic lifestyle and did not possess cranial adaptations for terrestrial feeding.     

Plastome organization and evolution of chloroplast genes in Cardamine species adapted to contrasting habitats

Background

Plastid genomes, also known as plastomes, are shaped by the selective forces acting on the fundamental cellular functions they code for and thus they are expected to preserve signatures of the adaptive path undertaken by different plant species during evolution. To identify molecular signatures of positive selection associated to adaptation to contrasting ecological niches, we sequenced with Solexa technology the plastomes of two congeneric Brassicaceae species with different habitat preference, Cardamine resedifolia and Cardamine impatiens.

Results

Following in-depth characterization of plastome organization, repeat patterns and gene space, the comparison of the newly sequenced plastomes between each other and with 15 fully sequenced Brassicaceae plastomes publically available in GenBank uncovered dynamic variation of the IR boundaries in the Cardamine lineage. We further detected signatures of positive selection in ten of the 75 protein-coding genes of the examined plastomes, identifying a range of chloroplast functions putatively involved in adaptive processes within the family. For instance, the three residues found to be under positive selection in RUBISCO could possibly be involved in the modulation of RUBISCO aggregation/activation and enzymatic specificty in Brassicaceae. In addition, our results points to differential evolutionary rates in Cardamine plastomes.

Conclusions

Overall our results support the existence of wider signatures of positive selection in the plastome of C. resedifolia, possibly as a consequence of adaptation to high altitude environments. We further provide a first characterization of the selective patterns shaping the Brassicaceae plastomes, which could help elucidate the driving forces underlying adaptation and evolution in this important plant family.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1498-0) contains supplementary material, which is available to authorized users.     

Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.