Morpho‐anatomical diversity of the underground systems of Arrojadoa (Cactaceae), an endemic Brazilian genus

Members of Arrojadoa exhibit a variety of underground structures that can originate from roots or stems. Although the development of underground structures of stem origin in Arrojadoa represents a unique trait among Cactaceae of eastern Brazil, no detailed reports on the morphological diversity of such structures are available. The present morpho‐anatomical study of the underground systems of Arrojadoa has demonstrated that a single species can exhibit one or more structural types, such as single or branched stem tubers, short thick stems and/or long subterranean stems, thick and fleshy contractile roots and normal fibrous roots. Various morpho‐anatomical structures relating to the underground storage systems in Arrojadoa spp. have also been observed, including thick contractile roots consisting mainly of secondary xylem formed by fibrous wood with wide‐band tracheids (WBTs) and underground stems with a wide cortical region and WBTs‐type wood. Based on the evidence presented, we suggest that such traits, together with the occurrence of contractile roots associated with underground stems, are important adaptive strategies for the survival of the plants during seasonal drought in areas of cerrado (savannah), campo rupestre (rocky uplands) and caatinga (dry thorny scrubland). © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 108–128.     

Seedling establishment,underground kinetics,and clonal reiteration: How do Potentilla inclinata and Inula ensifolia get their multifunctional subterranean systems?

The development of a seedling into an adult plant comprises various underground processes. Time-lapse photography (TLP) makes them visible. This is documented for Potentilla inclinata (Rosaceae) and Inula ensifolia (Asteraceae).After germination, P. inclinata develops a taproot system. Contraction phenomena pull the basal part of the shoot at least 10 mm into the soil. Later, several adventitious roots are generated, and thus the root system changes to a fibrous one. This is followed by cloning without separation of the ramets.Seedlings of I. ensifolia develop a weak primary root. At an early stage, adventitious roots are formed at the leaf rosette. This fibrous root system exerts a strong pulling effect on the shoot. After one vegetation period the basis of the rosette is approx. 30 mm under the soil surface. Cloning includes the formation of many new horizontal shoots, which conquer new sites.These two examples show three functional steps common in the developmental progress of subterranean systems: (I) establishment of the seedling, (II) innovation and survival of the young plant, and (III) reiteration (cloning and dispersal). However, to accomplish these basic development steps the diversity of subterranean systems is enhanced by different organographical components.Furthermore, the development of subterranean systems is a dynamic process consisting of two kinetic processes: the vertical movement during seedling establishment, which brings the innovation buds to a safe soil position, and the horizontal movement during dispersal, which conquers new sites.     

傅里叶变换红外光谱(FTIR)技术对滇龙胆组织培养的研究

野生药用植物资源的不断减少,使得寻找其原植物的合适替代品显得尤为重要。利用组培材料代替野生药用植物作为药源已取得重大进展,但利用傅里叶变换红外光谱(Fourier transform infrared spectroscopy,FTIR)技术筛选合适的组培材料作为野生药用植物替代资源方面的应用鲜有报道。本研究采用FTIR结合偏最小二乘判别分析(partial least squares discriminant analysis,PLS-DA)对滇龙胆组织培养形成的愈伤组织(肉质部、茎、叶)、增殖苗(肉质部、茎、叶)、生根苗(根、茎、叶)进行比较。结果显示:(1)从原始FTIR光谱图上看,滇龙胆肉质部和根部峰形相似,茎和叶峰形相似;(2)二阶导数光谱图扩大了样品间的差异。在龙胆苦苷的主要吸收峰1612 cm-1附近,吸收峰强度依次为:生根苗叶增殖苗叶和生根苗茎增殖苗茎愈伤组织叶,愈伤组织茎及肉质部、增殖苗肉质部和生根苗根部在该处无吸收峰;(3)PLS-DA得分图表明,同一组培阶段相同组织部位样品聚集在一起,而愈伤组织、增殖苗、生根苗及其各组织部位能够较好的分开。其中:肉质部、根部与茎叶之间距离较远,表明其化学成分和含量可能差异较大;肉质部和根部样品间距离较近,茎和叶样品间距离也较近。二阶导数光谱图显示,组培材料有望代替其原植物满足药用需求;若以龙胆苦苷含量为评价对象,生根苗叶则可能具有更大的开发潜能,有望代替野生滇龙胆以缓解其资源稀缺局面。本研究结果表明,采用傅里叶变换红外光谱法可以简便有效地对药用植物不同组培阶段不同组织部位的替代潜力及开发利用进行初步评估。     

THE NATURE OF THE FLESHY SHOOT OF ALLENROLFEA AND ALLIED GENERA

James , Lois E. (Whittier Coll., Whittier, Calif.), and Donald W. Kyhos . The nature of the fleshy shoot of Allenrolfea and allied genera. Amer. Jour. Bot. 48(2): 101–108. Illus. 1961.—A number of genera of the Chenopodiaceae are characterized by what have been called fleshy internodes which make them appear jointed. The fleshy tissue comprises on the average the outer ⅔ of what superficially appears to be the stem. This fleshy tissue contains a vascular network which is continuous with and similar to the vascular network in what has commonly been called the scalelike leaf. The problem is whether this fleshy vascularized tissue is the result of a downward prolongation of the base of the leaf or whether it is actually the true cortex of the stem. The present authors have found that Allenrolfea is uniquely suited to solving the problem of the so-called fleshy internodes of these allied genera. The origin and development of the fleshy tissue indicate it to be foliar. This conclusion is further substantiated by the shape of the fleshy tissue, its phyllotactic arrangement, the extent of the palisade tissue and underlying vascular network, the pattern of branching of the leaf traces, and the effect of secondary thickening and suberization.     

Morphology and development of the subterranean organs of the achlorophyllous Sciaphila polygyna (Triuridaceae)

The subterranean organs of the achlorophyllous Sciaphila polygyna (Triuridaceae) are described, depicted, and structurally explained for the first time. Unlike other Triuridaceae, the subterranean system of S. polygyna appears as a complex star-like structure of short, but thickened, roots as well as scale leaves and shoots. A complete series of sections revealed the following construction. In the axil of a scale leaf at a shoot of first order, a side shoot of second order as well as a pair of endogenous shoot-borne roots arise. This side shoot of second order also develops a scale leaf very early in ontogeny, which again gives rise to a side shoot of third order and a pair of shoot-borne roots. Other scale leaves at shoots of any order may also bear shoots and root pairs. This growth pattern occurs in a very close manner without internode elongation, resulting in the clumped, star-like appearance. The structures described superficially resemble the root systems of many mycoheterotrophic plants from other families. Comparisons with respect to how they develop, however, show that these similar root systems can result from distinct developmental patterns, suggesting independent evolutionary pathways and a considerable evolutionary pressure towards abbreviated and thickened roots in mycoheterotrophic plants. Possible advantages as well as evolutionary implications of the structures described are discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 295–301.     

Comparisons among biomass allocation and spatial distribution patterns of some vine,pteridophyte, and gymnosperm shoots

The interspecific allometry of leaf, stem, and reproductive biomass distal to stem diameter was determined for a total of 12 angiosperm vine, gymnosperm, and pteridophyte species to compare allocation patterns to vegetative and reproductive shoot organs. The allometry of stem diameter in terms of the distance from shoot apices also was determined to quantify the manner in which vines, gymnosperms, and pteridophyte stems tapered along their length. The stems of vine species were found to weigh more than those of arborescent gymnosperm species distal to any point of equivalent stem diameter. Vine species also distribute more of their stem mass to shoot length as opposed to girth than gymnosperm species. Vine stems also supported proportionally larger leaf and reproductive biomass in comparison to gymnosperm stems of equivalent diameter, yet partitioned their total shoot biomass more or less equally between leaf and stem biomass in the same manner as the gymnosperm species examined. The allometry of vine as well as gymnosperm leaf biomass with respect to stem biomass appeared to be slightly anisometric and negative, suggesting that more massive stems had proportionally less leaf biomass than their smaller, less massive counterparts. Vine stems could be approximated as very slender cones; the shape and geometry of gymnosperm stems complied with those of stubby, truncated cones whose top diameter (for those examined), on the average, equaled 28% of the basal diameter. In general terms, the interspecific allometry of vines was most similar to that of pteridophytes. Collectively, these data refute the commonly held notion that vine stems are simply more slender than those of species with self-supporting stems.     

Tissue localization of a submergence-induced 1-aminocyclopropane-1-carboxylic acid synthase in rice

At least two 1-aminocyclopropane-1-carboxylic acid synthase genes (ACS) are implicated in the submergence response of rice (Oryza sativa). Previously, the OS-ACS5 gene has been shown to be induced during short- as well as long-term complete submergence of seedlings and to be controlled by a balance of gibberellin and abscisic acid in both lowland and deepwater rice. This study demonstrates that OS-ACS5 mRNA is localized in specific tissues and cells both during normal development and in response to complete submergence. The temporal and spatial regulation of OS-ACS5 expression is presented by in situ hybridization and histochemical analysis of beta-glucuronidase (GUS) activity in transgenic rice carrying an OS-ACS5-gus fusion. Whole-mount in situ hybridization revealed that in air-grown rice seedlings, OS-ACS5 was expressed at a low level in the shoot apex, meristems, leaf, and adventitious root primordia, and in vascular tissues of nonelongated stems and leaf sheaths. In response to complete submergence, the expression in vascular bundles of young stems and leaf sheaths was strongly induced. The results of histochemical GUS assays were consistent with those found by whole-mount in situ hybridization. Our findings suggest that OS-ACS5 plays a role in vegetative growth of rice under normal conditions and is also recruited for enhanced growth upon complete submergence. The possible implication of OS-ACS5 in root-shoot communication during submergence stress and its putative role in aerenchyma formation upon low-oxygen stress are discussed.     

Adventitious shoot formation in decapitated dicotyledonous seedlings starts with regeneration of abnormal leaves from cells not located in a shoot apical meristem

Regeneration of new shoots in plant tissue culture is often associated with appearance of abnormally shaped leaves. We used the adventitious shoot regeneration response induced by decapitation (removal of all preformed shoot apical meristems, leaving a single cotyledon) of greenhouse-grown cotyledon-stage seedlings to test the hypothesis that such abnormal leaf formation is a normal regeneration progression following wounding and is not conditioned by tissue culture. To understand why shoot regeneration starts with defective organogenesis, the regeneration response was characterized by morphology and scanning electron and light microscopy in decapitated cotyledon-stage Cucurbita pepo seedlings. Several leaf primordia were observed to regenerate prior to differentiation of a de novo shoot apical meristem from dividing cells on the wound surface. Early regenerating primordia have a greatly distorted structure with dramatically altered dorsoventrality. Aberrant leaf morphogenesis in C. pepo gradually disappears as leaves eventually originate from a de novo adventitious shoot apical meristem, recovering normal phyllotaxis. Similarly, following comparable decapitation of seedlings from a number of families (Chenopodiaceae, Compositae, Convolvulaceae, Cucurbitaceae, Cruciferae, Fabaceae, Malvaceae, Papaveraceae, and Solanaceae) of several dicotyledonous clades (Ranunculales, Caryophyllales, Asterids, and Rosids), stems are regenerated bearing abnormal leaves; the normal leaf shape is gradually recovered. Some of the transient leaf developmental defects observed are similar to responses to mutations in leaf shape or shoot apical meristem function. Many species temporarily express this leaf development pathway, which is manifest in exceptional circumstances such as during recovery from excision of all preformed shoot meristems of a seedling.     

Life History Evolution in Amphicarpic Plants

Abstract Plants with dimorphic flowers or seeds provide excellent material for the study of life history evolution because the dimorphism often involves measurable differences in morphology, size, number, or genetic relatedness. For amphicarpic plants, the proportion of aerial: subterranean morphs produced is highly variable (from 0 to > 100) and related to both environmental and genetic factors. Plants from aerial seeds produce lower ratios of aerial: subterranean morphs than those from subterranean seeds. Despite substantial variation, subterranean seeds are generally heavier than aerial seeds (but fewer) and produce vigorous seedlings with high survivorship and high fitness. Adaptive advantages of subterranean seeds include retention of offspring in favorable parental microhabitats, protection of seeds from herbivory, predation, or fire, and avoidance of desiccating conditions on the soil surface; potential disadvantages include lack of gene exchange, high energy costs, limited dispersal, and sibling competition. For the few species studied, aerial reproduction is more plastic than subterranean reproduction and more likely to be affected by environmental conditions. Quantitative genetic analyses of a population of the annual grass Amphicarpum purshii have revealed lower heritabilites for subterranean relative to aerial reproductive traits. Subterranean seed number and mass show genetic correlations with shoot mass while aerial seed number and mass do not; seed set percentages of both seed types as well as percentage allocation to both reproductive morphs show negative genetic correlations with shoot mass. In this Amphicarpum population, directional selection on shoot mass may indirectly select for increased subterranean (but not aerial) seed output.     

Phylloclade development in the Asparagaceae: an example of homoeosis

Phylloclades are traditionally defined as flattened, determinate, leaf-like stems primarily on the basis of their axillary position. However, because the literature is replete with controversy over the morphological interpretation of these organs, a study of phylloclade development in comparison with leaf and stem development was undertaken in four closely related species of the Asparagaceae: Ruscus aculeatus, Danae racemosa, Semele androgyna and Asparagus densiflorus. Results reveal a continuum in phylloclade development from very leaf-like forms, such as those of Danae , via the more intermediate types of Ruscus , to the gradually more shootlike forms of Semele and Asparagus. This continuum results from a differential expression of stem (or shoot) and leaf characteristics in an axillary position. When stem (or shoot) and leaf features are combined, as in the fertile phylloclade of Ruscus , an intermediate organ is formed. Phylloclades are a form of evolutionary novelty that exemplifies the phenomenon of homoeosis, which is the transference of features from one organ to another. Developmentally, this means that leaf features are expressed by the axillary meristem.     

不同光质光周期对樱桃萝卜生长发育及营养品质的影响

光质和光周期可以调节植物生长过程中肉质根等地下贮藏器官的形成。该试验以‘昆优2号’樱桃萝卜为试验材料,以白光(W)为对照,研究8R2B、5R5B、2R8B红蓝光配比光质(红蓝光比例分别为8∶2、5∶5、2∶8)与8L/16D、12L/12D、16L/8D光周期(光照/黑暗分别为8 h/16 h、12 h/12 h、16 h/8 h)组合对樱桃萝卜生长发育和营养品质的影响,筛选适合樱桃萝卜生长的光质光周期组合。结果表明:长光照(16L/8D)W、8R2B处理对樱桃萝卜叶片生长、肉质根大小、根冠比都有显著的促进作用,长光照W有利于樱桃萝卜更快形成膨大的肉质根;长光照W、8R2B处理下的樱桃萝卜肉质根多酚含量、抗氧化能力差异不大;长光照W利于樱桃萝卜肉质根可溶性糖、可溶性蛋白积累,长光照8R2B有利于肉质根可溶性蛋白积累。研究发现,适当的光质和光周期组合可显著促进樱桃萝卜叶片生长、肉质根膨大以及根中营养物质的积累,从而获得良好的长势和营养品质,并以长光照(16L/8D即光照/黑暗为16 h/8 h)白光组合最佳。     

吊丝单竹笋期叶片特性研究

对3种不同年龄的吊丝单竹株在不同发笋时期、不同竹冠部位的叶片比叶重、光合、呼吸性状及N、P、K含量等进行测定,结果表明:吊丝单竹在不同发笋时期、不同竹冠部位各项指标均有较大差异。比叶重发笋盛期高于初期和末期,随着竹株年龄增大而减小,随着竹冠的升高而增大;净光合速率盛期高于初期,光呼吸速率和暗呼吸速率则初期高于盛期,并且不同竹龄、不同竹冠部位之间均表现出较大的差异;N、P、K含量由发笋初期至盛期逐渐减小,N含量以1a>2a>3a,以竹冠中部最高,P、K含量则随着竹冠的升高而减小,随着竹株年龄的增大而增大。这为经营管理吊丝单竹林提供科学依据。     

荸荠营养器官的发育与解剖学研究

荸荠同化茎起源于肉质主茎倒２或倒３叶的叶腋内。同化茎基部着生二鞘状叶,鞘状叶对早期同化茎穿出土面具保护作用。匍匐茎大多起源于同化茎基部鞘状叶的叶腋内。当植株开始抽生花茎时,地下匍匐茎顶端开始膨大。球茎的膨大是匍匐茎顶端５－６节的基本组织经细胞有丝分裂,增加细胞数目,然后由细胞体积的扩大来实现的。球茎具足够的营养物质供来年顶芽萌发的需要,故属水生植物冬芽的性质。     

The Mechanical Roles of Clasping Leaf Sheaths: Evidence fromArundinaria tecta(Poaceae) Shoots Subjected to Bending and Twisting Forces

Representative shoot segments of the grass speciesArundinariatéctaconsisting of one intact internode and its subtendingnode and clasping leaf sheath were tested to determine the mechanicalinfluence of the leaf sheath on the ability of stems to resistbending and twisting forces. These segments were also used tomeasure shoot morphometry and composite tissue Young's and shearmoduli (EandG,respectively) to simulate the global deformationpatterns attending bending and twisting by means of finite elementanalyses. On average, leaf sheaths contributed 33% of the overallbending stiffness and 43% of the overall torsional stiffnessof stem segments. Comparisons betweenEandGof isolated internodesand leaf sheaths indicated that sheaths were composed of stiffertissues measured either in bending or twisting. Thus, leaf sheathscould act as an external cylindrical brace composed of stiffermaterials than those of the internodes they enveloped. The magnitudesof internodalEandGwere correlated with internodal shape suchthat the ability of internodes to resist twisting relative tothe ability to resist bending forces decreased as internodesbecame more slender or developed thinner walls (both of whichoccur in an acropetal direction from the base to the tip ofshoots). Finite element simulations predicted that, in bending,the leaf sheath laterally braces internodal walls as they tendto ovalize in cross section and push against its inner surfacewhich ovalizes to a lesser extent in the plane normal to thecurvature of shoot flexure. In twisting, the successive ovalizedtransections of internodal walls assumed a helical pattern alongthe length of shoot segments. This helical deformation patternwas attended by an inner lateral contraction of internodal wallsthat was less developed in the leaf sheath that thus provideddecreasing mechanical support to the internode as the lateralcontraction of internodal walls amplified. The twisting of internodesand sheaths was also predicted to concentrate tensile and shearstrains in the nodal diaphragm. Here stress intensities sufficientto produce tissue shear failure were concentrated at two opposingpoints on the surface of the diaphragm. Finite element analysesthus identified a potential weak point in the mechanical constructionof hollow, septate shoots that are, nevertheless, more thanadequately stiff to support their own weight, yet sufficientlyflexible to twist without irreparable damage in normal winds.Copyright1998 Annals of Botany Company Plant stems; nodes; internodes; leaf sheaths; elastic moduli; wind lodging; biomechanics.     

Phylogenetic diversification of Equisetum (Equisetales) as inferred from Lower Cretaceous species of British Columbia, Canada

Three types of anatomically preserved vegetative shoots with features that characterize crown group Equisetum have been discovered in Lower Cretaceous deposits (≈136 Ma) of British Columbia, Canada, suggesting the genus is much older than currently believed. Specimens include two types of aerial shoots described as E. haukeanum sp. nov. and E. vancouverense sp. nov. and one type of subterranean rhizome. Shoots are 1-2 mm in diameter, jointed, and in cross section have fluted stems with a hollow pith. Distinctive patterns of cortical sclerenchyma and different ridge morphologies characterize each shoot morphotype. Nodes display irregular branching, highly fused leaf sheaths, and a nodal diaphragm. The aerial stem morphospecies have vallecular canals on alternating radii with carinal canals of an equisetostele surrounded by only a few tracheids. No secondary tissues are produced. Bands of surficial stomata flank the furrows of one morphospecies. Rhizomes and aerial shoots are of a similar size, suggesting that the plants were equivalent in stature to the smallest living Equisetum species. These fossils augment our understanding of evolutionary transformations that led from Paleozoic Archaeocalamitaceae and Calamitaceae to crown group Equisetaceae, suggesting that the initial diversification of Equisetum began far earlier than suggested by molecular-clock-based estimates.     

THE PHYSIOLOGICAL BASIS OF MORPHOLOGICAL POLARITY IN REGENERATION. I

1. In Bryophyllum calycinum two apical leaves suppress the shoot formation in all the dormant buds situated basally from the leaf; one apical leaf suppresses the shoot formation in the basal buds situated in the same half of the stem where the leaf is, and, if one-half of the petiole of such a leaf is removed, the growth of basal buds in one quadrant of the stem is suppressed. 2. This inhibitory influence of a leaf upon shoot formation in the basal part of a stem is diminished or disappears when the mass of the leaf is reduced below a certain limit. 3. The inhibitory influence of an apical leaf upon the growth of shoots in horizontally suspended stems is greater when the leaf is on the upper than when it is on the lower side of the stem. 4. All these facts suggest the possibility that the inhibitory influence of the leaf upon shoot formation is due to inhibitory substances secreted by the leaf and carried by the sap from the leaf towards the base of the stem. 5. An apical leaf accelerates root formation in the basal part of a stem and this accelerating effect increases with the mass of the leaf. 6. This inhibitory influence of a leaf upon shoot formation and the favoring influence upon root formation in the more basally situated parts of the stem is one of the factors determining the polar character of regeneration.     

Tissue and cellular phosphorus storage during development of phosphorus toxicity in Hakea prostrata (Proteaceae)

Storage of phosphorus (P) in stem tissue is important in Mediterranean Proteaceae, because proteoid root growth and P uptake is greatest during winter, whereas shoot growth occurs mostly in summer. This has prompted the present investigation of the P distribution amongst roots, stems, and leaves of Hakea prostrata R.Br. (Proteaceae) when grown in nutrient solutions at ten P-supply rates. Glasshouse experiments were carried out during both winter and summer months. For plants grown in the low-P range (0, 0.3, 1.2, 3.0, or 6.0 micromol d(-1)) the root [P] was > stem and leaf [P]. In contrast, leaf [P] > stem and root [P] for plants grown in the high-P range (6.0, 30, 60, 150, or 300 micromol P d(-1)). At the highest P-supply rates, the capacity for P storage in stems and roots appears to have been exceeded, and leaf [P] thereafter increased dramatically to approximately 10 mg P g(-1) dry mass. This high leaf [P] was coincident with foliar symptoms of P toxicity which were similar to those described for many other species, including non-Proteaceae. The published values (tissue [P]) at which P toxicity occurs in a range of species are summarized. X-ray microanalysis of frozen, full-hydrated leaves revealed that the [P] in vacuoles of epidermal, palisade and bundle-sheath cells were in the mM range when plants were grown at low P-supply, even though very low leaf [P] was measured in bulk leaf samples. At higher P-supply rates, P accumulated in vacuoles of palisade cells which were associated with decreased photosynthetic rates.     

A NEW SPECIES OF MAIANTHEMUM (LILIACEAE) FROM COSTA RICA WITH AN UPRIGHT AND AERIAL RHIZOME

A new species of Maianthemum (Liliaceae) was discovered in a tropical montane bog at an elevation of 2,500 m in central Costa Rica. The leaf bearing stems and inflorescences are supported by a stiff, erect, branching sympodium that consists of the persistent lower portions of older shoots. This lower portion corresponds in morphology and anatomy to the horizontal subterranean rhizome that is characteristic of all previously described species of Maianthemum. An erect woody habit is rare in the Lily family; an upright aerial rhizome is unique within the genus.     

Shoot meristem function and leaf polarity: the role of class III HD-ZIP genes

The shoot apical meristem comprises an organized cluster of cells with a central region population of self-maintaining stem cells providing peripheral region cells that are recruited to form differentiated lateral organs. Leaves, the principal lateral organ of the shoot, develop as polar structures typically with distinct dorsoventrality. Interdependent interactions between the meristem and developing leaf provide essential cues that serve both to maintain the meristem and to pattern dorsoventrality in the initiating leaf. A key component of both processes are the class III HD-ZIP genes. Current findings are defining the developmental role of members of this family and are identifying multiple mechanisms controlling expression of these genes.