Derived woodiness and annual habit evolved in African umbellifers as alternative solutions for coping with drought.

BACKGROUND: One of the major trends in angiosperm evolution was the shift from woody to herbaceous habit. However, reversals known as derived woodiness have also been reported in numerous, distantly related clades. Among theories evoked to explain the factors promoting the evolution of derived woodiness are moderate climate theory and cavitation theory. The first assumes that woody habit evolves in response to mild climate allowing for prolonged life span, which in turn leads to bigger and woodier bodies. The second sees woodiness as a result of natural selection for higher cavitation resistance in seasonally dry environments. Here, we compare climatic niches of woody and herbaceous, mostly southern African, umbellifers from the Lefebvrea clade to assess whether woody taxa in fact occur in markedly drier habitats. We also calibrate their phylogeny to estimate when derived woodiness evolved. Finally, we describe the wood anatomy of selected woody and herbaceous taxa to see if life forms are linked to any particular wood traits. RESULTS: The evolution of derived woodiness in chamaephytes and phanerophytes as well as the shifts to short-lived annual therophytes in the Lefebvrea clade took place at roughly the same time: in the Late Miocene during a trend of global climate aridification. Climatic niches of woody and herbaceous genera from the Cape Floristic Region overlap. There are only two genera with distinctly different climatic preferences: they are herbaceous and occur outside of the Cape Floristic Region. Therefore, studied herbs have an overall climatic niche wider than their woody cousins. Woody and herbaceous species do not differ in qualitative wood anatomy, which is more affected by stem architecture and, probably, reproductive strategy than by habit. CONCLUSIONS: Palaeodrought was likely a stimulus for the evolution of derived woodiness in the Lefebvrea clade, supporting the cavitation theory. The concurrent evolution of short-lived annuals withering before summer exemplifies an alternative solution to the same problem of drought-induced cavitation. Changes of the life form were most likely neither spurred nor precluded by any qualitative wood traits, which in turn are more affected by internode length and probably also reproductive strategy.

Investigating Schizocarp morphology as a taxonomic tool in study of Apiaceae family by utilizing LM and SEM techniques.

In present study, the schizocarp morphology of 14 species belonging to Apiaceae family has been investigated. Light microscopy (LM) and scanning electron microscopy (SEM) have been utilized to highlight qualitative and quantitative features of studied species. Variations have been observed in macro- and micro-morphological features such as color, shape, symmetry, length, width, apex, epicuticular projections, surface patterns, anticlinal, and periclinal wall patterns. Schizocarp shapes observed were oval, round, triangular, linear, elliptic, and globose. Fruit was either homomorphic or heteromorphic. Crystalloids, stellate hair, multicellular spines, and platelets were mostly observed epicuticular projections. Surface patterns on the fruit surface were striate, rugulate-striate, reticulate, and striato-knotted. Both macro- and micro-morphological characters can serve as an important tool in classifying Apiaceae family at various taxonomic ranks. Substantial variations observed can assist as useful constraints at various taxonomic levels as they provide reliable and constant details. Disparities observed in schizocarp features can pave a path for Apiaceae family classification based on phylogenetic and molecular studies.

Specific features of root aerenchyma formation in Sium latifolium L. (Apiaceae).

Types of aerenchyma formation in adventive roots of wild plant S. latifolium L. are described for the first time. Specific cell divisions and patterns of cell growth in the inner and outer root cortex at the initial stages of aerenchyma formation are highlighted in detail. Destructive processes in cells occurred under aerenchyma formation are considered in the view of known stages of programmed cell death in plants.

Easy Come, Easy Go: Capillary Forces Enable Rapid Refilling of Embolized Primary Xylem Vessels.

Protoxylem plays an important role in the hydraulic function of vascular systems of both herbaceous and woody plants, but relatively little is known about the processes underlying the maintenance of protoxylem function in long-lived tissues. In this study, embolism repair was investigated in relation to xylem structure in two cushion plant species, Azorella macquariensis and Colobanthus muscoides, in which vascular water transport depends on protoxylem. Their protoxylem vessels consisted of a primary wall with helical thickenings that effectively formed a pit channel, with the primary wall being the pit channel membrane. Stem protoxylem was organized such that the pit channel membranes connected vessels with paratracheal parenchyma or other protoxylem vessels and were not exposed directly to air spaces. Embolism was experimentally induced in excised vascular tissue and detached shoots by exposing them briefly to air. When water was resupplied, embolized vessels refilled within tens of seconds (excised tissue) to a few minutes (detached shoots) with water sourced from either adjacent parenchyma or water-filled vessels. Refilling occurred in two phases: (1) water refilled xylem pit channels, simplifying bubble shape to a rod with two menisci; and (2) the bubble contracted as the resorption front advanced, dissolving air along the way. Physical properties of the protoxylem vessels (namely pit channel membrane porosity, hydrophilic walls, vessel dimensions, and helical thickenings) promoted rapid refilling of embolized conduits independent of root pressure. These results have implications for the maintenance of vascular function in both herbaceous and woody species, because protoxylem plays a major role in the hydraulic systems of leaves, elongating stems, and roots.

Morphometrics of Daucus (Apiaceae): a counterpart to a phylogenomic study.

PREMISE OF STUDY: Molecular phylogenetics of genome-scale data sets (phylogenomics) often produces phylogenetic trees with unprecedented resolution. A companion phylogenomics analysis of Daucus using 94 conserved nuclear orthologs supported many of the traditional species but showed unexpected results that require morphological analyses to help interpret them in a practical taxonomic context. METHODS: We evaluated character state distributions, stepwise discriminant analyses, canonical variate analyses, and hierarchical cluster analyses from 40 morphological characters from 81 accessions of 14 taxa of Daucus and eight species in related genera in an experimental plot. KEY RESULTS: Most characters showed tremendous variation with character state overlap across many taxa. Multivariate analyses separated the outgroup taxa easily from the Daucus ingroup. Concordant with molecular analyses, most species form phenetic groups, except the same taxa that are problematical in the molecular results: (1) the subspecies of D. carota, (2) D. sahariensis and D. syrticus, and (3) D. broteri and D. guttatus. CONCLUSIONS: Phenetic analyses, in combination with molecular data, support many Daucus species, but mostly by overlapping ranges of size and meristic variation. The subspecies of D. carota are poorly separated morphologically, are paraphyletic, and all could be recognized at the subspecies rank under D. carota. Daucus sahariensis and D. syrticus are so similar morphologically that they could be placed in synonymy. Combined molecular and morphological data support three species in accessions previously identified as D. broteri and D. guttatus. Molecular and morphological results support the new combination Daucus carota subsp. capillifolius.

Andromonoecy and developmental plasticity in Chaerophyllum bulbosum (Apiaceae-Apioideae).

BACKGROUND AND AIMS: Andromonoecy, the presence of hermaphrodite and male flowers in the same individual, is genetically fixed or induced, e.g. by fruit set. Little is known about the forces triggering andromonoecy in the Apiaceae. In the present study, a natural population of the protandrous Chaerophyllum bulbosum was investigated to elucidate architectural constraints and effects of resource reallocation. METHODS: Three sets of plants (each n = 15) were treated by hand pollination, pollinator exclusion and removal of low-order inflorescences. Fifteen untreated plants were left as controls. KEY RESULTS: Untreated plants produce umbels up to the third branch order, with increasing proportions of male flowers from 15 % (terminal umbel) to 100 % (third-order umbels). Fruit set correspondingly decreases from 70% (terminal umbel) to <10 % (second-order umbels). Insignificant differences from hand-pollinated plants do not reveal any sign of pollinator limitation at the study site. Bagged individuals show the same increase in male flowers with age as untreated plants, indicating that the presence of andromonoecy is not induced by fruit set. After umbel removal, individuals tend to present a higher number of hermaphrodite flowers and fruits in the umbels of second and third order. Three plants (25 %) produced an additional branch order composed of 100 % male umbels. CONCLUSIONS: Inherited andromonoecy and the plastic response to environmental conditions are interpreted as a self-regulating system saving investment costs and optimizing fruit set at the same time.

[Study on macroscopic and microscopic identification of Saposhnikovia divaricata and its counterfeits].

OBJECTIVE: To provide an identification method for the roots of Saposhnikovia divaricata and its three counterfeits. METHODS: Macroscopic identification and microscopic identification of root transverse section and powder were carried out to distinguish these four species. RESULTS: For macroscopic characteristics, Saposhnikoviae Radix and its counterfeits can be distinguished by the head of the residual leaf and sections. As for microscopic identification, the feature was not obvious. But there were some differences to distinguish them,such as the number of cork layer, cambium was evident or not, the number of the xylem catheter,the presence or absence of large oil pipe and longitudinal cracks between the part from cortex to xylem. CONCLUSION: This is a simple and accurate method for distinguish Saposhnikoviae Radix and its counterfeits.

The phylogenetic significance of the carpophore in Apiaceae.

BACKGROUND AND AIMS: Fruit structural characters have traditionally been important in the taxonomy of the family Apiaceae. Previous investigations using a limited number of taxa have shown that the carpophore may be especially useful in helping to circumscribe subfamily Azorelloideae. The present study examines, for the first time, carpophore structure in 92 species from 43 genera, representing all subfamilies of Apiaceae, and including all genera assigned to subfamily Azorelloideae. Phylogenetic interpretations are made for the first time, using all available information, and a standard terminology is proposed to describe the various character states found in carpophores. METHODS: Carpophore structure was studied in detail using light microscopy. KEY RESULTS: Carpophores, when present, may be categorized into two main groups (B and C) based mainly on the arrangement of the vascular bundles in transverse section, and further divided into six sub-types according to the length of the carpophore (short in B1 and C1) and whether they are entire (B1-B3 and C1) or bifurcate (B4 and C2). Free carpophores are absent in subfamily Mackinlayoideae, and in tribes Lichtensteinieae and Phlyctidocarpeae, which have two opposite vascular bundles (Group A). Entire carpophores with one or two vascular bundles, or bifurcate carpophores with lateral vascular bundles (arranged side by side within the commissural plane), are the main types characterizing Azorelloideae. The short, hygroscopic carpophores found in Choritaenia are unique in Apiaceae and provide additional evidence for the exclusion of this genus from Azorelloideae. Carpophore type C2 is typical for most Apioideae sensu lato (exceptions are, for example, Arctopus and Alepidea, which have type B2). CONCLUSIONS: A single carpophore and ventral vascular bundles not forming free carpophores are proposed to be the ancestral conditions in Apiaceae, while bifurcate carpophores with opposite vascular bundles are the derived state, present in most Apioideae. Secondary reductions seem to have occurred in several unrelated lineages in all major groups, e.g. many Azorelloideae, several protoapioids (including nearly all members of the tribe Saniculeae) and 29 euapioid genera (e.g. some Oenantheae).

[Comparison on botanical morphology in different populations of Changium smyrnioides].

OBJECTIVE: To analyse botanical morphology in different populations and give a reference for germplasm evaluation of Changium smyrnioides. METHOD: Roots and leaves were comprehensively compared among 10 populations of Ch. smyrnioides in the main distribution area, the foliar morphology similarity coefficient was calculated, and the systematic relationships were constructed based on UPGMA method. RESULT: There was significant morphological difference among populations of Ch. smyrnioides. Plant shape was divided into the loosed-type, compacted-type and intermediated-type. Leaf split was divided into the lobated-type, parted-type and intermediate-type. Root shape was divided into spherical-type, column-type and spindle-type. Foliar morphology similarity coefficient differed significantly and ranged from 0.234 1 to 0.985 1 among 10 populations of Ch. smyrnioides. Zijinshan population was divided into a unique group when similarity coefficient was 0.37. CONCLUSION: Plant tightness, lobation depth and root shape could be used as the basis for germplasm evaluation on Ch. smyrnioides. Zijinshan population could be used as a special germplasm to be exploited.

[Difference of shapes and propertiesand microscopic frameworks between wild and cultivated Radix Saposhnikovia].

OBJECTIVE: To find the difference of the shapes and properties and the microscopic frameworks between wild and cultivated Radix Saposhnikovia. METHOD: The shapes and properties, the characters of transverse section, the powder and disintegrated tissue of roots of medical materials were compared by microscopic measuring. RESULT: Wild Radix Saposhnikovia had a long conical or cylindrical root, and fewer root branches. It showed a close annulus grain on top root, cortical section of root in light brown colour, many brown oil spots and possessed typical odor, While cultivated Radix Saposhnikovia had many root branches, and showed less annulus grain on top root, cortical section of root in light yellow brown colour, less brown oil spots and possessed light odor. The difference of microscopic histological structure was that wild Radix Saposhnikovia had phloem transverse section of root with many rotundity oil tube lining up 10-22 rings, xylem vessel with radiate rank, and indistinct annual ring. While cultivated Radix Saposhnikovia had phloem transverse section of root with oil tube lining up 10-11 rings and xylem vessel with distinct annual ring. CONCLUSION: There exists several differences between wild and cultivated Radix Saposhnikovia in shapes and properties and differences of microscopic frameworks. The main characteristics are the differences of shapes and numbers of oil tube of phloem transverse section of root. The cultivated Radix Saposhnikovia of 1-4 years can be recognized by annual rings of xylem vessel.

Characterization of compounds from the roots of Saposhnikovia divaricata by high-performance liquid chromatography coupled with electrospray ionization tandem mass spectrometry.

In this study, four types of compounds including coumarins, chromones, furoylmethyl amino acid derivative and benzofuran glycoside were isolated from the roots of Saposhnikovia divaricata. The electrospray ionization (ESI) mass spectral fragmentation pathways of these compounds were proposed. In particular, the ESI-MS(n) fragmentation behavior of linear dihydrofurocoumarins, dihydrofuro- and dihydropyranochromones were deduced in detail. For the linear dihydrofurocoumarins, the fragmentation was triggered by the initial loss of the C-4' substituting group. Then, the characteristic ions were observed followed by the losses of 15, 18, 28 and 46 Da. It is noteworthy that the elimination of H(2)O (18 Da) from the cleavage of the dihydrofuran ring is reported for the first time. For the linear dihydrofurochromones, characteristic eliminations of 18, 48 and 72 Da were observed. The loss of 18 Da could arise from two different fragmentation pathways, and the observed ion was composed of a mixture of two different structural ions. For the linear dihydropyranochromones, it was found that the dihydropyran ring was converted into the pyran ring by the elimination of the C-3' substituting group. This fragmentation was followed by the diagnostic losses of 18, 28, 42 and 54 Da in tandem mass spectrometry. The above fragmentation rules were successfully applied for the analysis of the chemical constituents of the roots of Saposhnikovia divaricata. A total of 32 compounds were identified or tentatively characterized by HPLC/DAD/ESI-MS(n). Among them, eight compounds were new and seven compounds were reported from that genus for the first time.

Adaptations to increasing hydraulic stress: morphology, hydrodynamics and fitness of two higher aquatic plant species.

Sessile organisms often exhibit morphological changes in response to permanent exposure to mechanical stimulation (wind or water movements). The adaptive value of these morphological changes (hydrodynamic performance and consequences on fitness) has not been studied extensively, particularly for higher plants submitted to flow stress. The aim was to determine the adaptive value of morphological patterns observed within two higher aquatic plant species, Berula erecta and Mentha aquatica, growing along a natural flow stress gradient. The hydrodynamic ability of each ramet was investigated through quantitative variables (drag coefficient and E-value). Fitness-related traits based on vegetative growth and clonal multiplication were assessed for each individual. For both species, the drag coefficient and the E-value were explained only to a limited extent by the morphological traits used. B. erecta exhibited a reduction in size and low overall plant drag at higher flow velocities, despite high drag values relative to leaf area, due to a low flexibility. The plants maintained their fitness, at least in part, through biomass reallocation: one tall ramet at low velocity, but shorter individuals with many interconnected stolons when flow velocity increased. For M. aquatica, morphological differences along the velocity gradient did not lead to greater hydrodynamic performance. Plant size increased with increasing velocities, suggesting the indirect effects of current favouring growth in high velocities. The fitness-related traits did not demonstrate lower plant fitness for high velocities. Different developmental constraints linked to plant morphology and trade-offs between major plant functions probably lead to different plant responses to flow stress.

Comparative studies on phenotypic plasticity of two herbs, Changium smyrnioides and Anthriscus sylvestris.

The endangered medicinal herb, Changium smyrnioides can only be found in deciduous forest gaps within the middle to northern subtropical broad-leaved evergreen forest zone of China. The considerable plasticity of its shoot and root structure helps it to capture light more effectively in winter and early spring, and to adapt to the soil moisture conditions in its narrow habitat. Another medicinal plant, Anthriscus sylvestris, is of similar economic importance but commonly dis-tributed widely. In contrast to C. smyrnioides, it has low structural plasticity. It is also specialized to adapt to the moist and sunny environment, where habitat, such as the banks of creeks and rivers, is abundant.

[Pharmacognostical studies on radix Glehniae (Glehnia littoralis)].

OBJECT: To identify the roots of Glehnia littoralis Fr. Schmidt ex Miq., and compare the chemical constituents of the root skin and the roots with no skin. METHODS: The roots were identified by morphological and microscopic identification and TLC. RESULTS: The characteristics of the secretory canal, ray and starch grain can be used to identify the histology and powder of the roots. The chemical constituents of the root skin and the roots with no skin are similar. CONCLUSION: The characteristics of the morphology, histology and powder can be used to identify the roots of Glehnia littoralis (Radix Glehniae).

[Microscopic and TLC identification on the fruits of ten species plants for Umbelliferae].

The fruits of ten species plants of Umbelliferae, including the fruits of Peucedanum decursiyum, Saposhnikovia divaricata, Peucedanum terebinthaceun, Anethum graveolens, Cnidium monnieri, Angelica sinensis, Foeniculum vulgate, Angelica polymorpha, Ferula tunnshanica and Cicuta virosa were identified on histology and TLC.

Pithiness in plants: I. The effect of mechanical perturbation and the involvement of ethylene in petiole pithiness in celery.

Mechanical perturbation (MP) applied to celery (Appium graveolens L. cv. Florida 683) leaf petioles or ethephon application to the plant did not induce thigmomorphogenesis (inhibition of elongation and increase in thickness of the petiole). However, the two treatments did cause the parenchyma breakdown which leads to pithiness or increased natural pithiness, mainly at the base of the petiole. Nevertheless, MP (but not ethephon) decreased the severity of drought-stress or GA3-induced pithiness. Although MP stimulates ethylene production, mainly at the middle part of the petiole, it seems that the protection by MP of the petiole may not be directly mediated by ethylene production. The exposure of the plant to drought stress brought about an increase in ethylene evolution. Upon reirrigating the plants, the first steps of pithiness were accompanied by a sharp decline in ethylene production. This decrease might be due to membrane disruption. The increase in ethylene production during drought stress may be one of the events which stimulate pithiness of the celery leaf petiole.