Primary and secondary thickening in the stem of Cordyline fruticosa (Agavaceae)

The growth in thickness of monocotyledon stems can be either primary, or primary and secondary. Most of the authors consider this thickening as a result of the PTM (Primary Thickening Meristem) and the STM (Secondary Thickening Meristem) activity. There are differences in the interpretation of which meristem would be responsible for primary thickening. In Cordyline fruticosa the procambium forms two types of vascular bundles: collateral leaf traces (with proto and metaxylem and proto and metaphloem), and concentric cauline bundles (with metaxylem and metaphloem). The procambium also forms the pericycle, the outermost layer of the vascular cylinder consisting of smaller and less intensely colored cells that are divided irregularly to form new vascular bundles. The pericycle continues the procambial activity, but only produces concentric cauline bundles. It was possible to conclude that the pericycle is responsible for the primary thickening of this species. Further away from the apex, the pericyclic cells undergo periclinal divisions and produce a meristematic layer: the secondary thickening meristem. The analysis of serial sections shows that the pericycle and STM are continuous in this species, and it is clear that the STM originates in the pericycle.The endodermis is acknowledged only as the innermost layer of the cortex.

Primary and secondary development of Cyperus giganteus Vahl rhizome (Cyperaceae)

In Cyperus giganteus, like in other Monocotyledoneae, the protoderm, procambium, fundamental meristem and primary thickening meristem (PTM) are differentiated from the rhizome promeristem. The PTM produces the inner cortical parenchyma, endodermis, pericycle and amphivasal vascular bundles, which are formed by the procambium too. After the primary body differentiates, cellular divisions continue only in the pericycle, and originate an irregular vascular system with vessel elements shorter and more branched than those found in the primary growth. This change of activity in the pericycle defines a secondary growth, where the secondary thickening meristem (STM) is the pericycle itself.

Diferenciação dos feixes vasculares e dos elementos traqueais no rizoma de algumas Cyperaceae

Foram analisados os rizomas de Bulbostylis paradoxa Ness, Cyperus giganteus Vahl, C. odoratus L., Fuirena umbellata Rottb. e Hypolytrum schraderianum Ness. O corpo primário é resultante da atividade dos meristemas apicais e do meristema de espessamento primário (MEP). Também ocorre crescimento em espessura, que é decorrente da atividade do meristema de espessamento secundário (MES). O procâmbio e o MEP originam feixes colaterais em H. schraderianum e feixes anfivasais nas demais espécies. Entretanto, todos os feixes que têm protofloema e protoxilema são de origem procambial. O MES produz floema e xilema constituindo um tecido vascular único. Elementos de vaso foram encontrados na maioria dos caules em estrutura primária e secundária, com exceção de H. schraderianum que, na estrutura secundária, contém apenas traqueídes, informação que respalda a ocorrência de crescimento secundário nas Cyperaceae. Os elementos de vaso apresentam grande variação morfológica; em estrutura primária, geralmente são mais alongados, com apêndices. Os elementos de vaso do crescimento secundário são relativamente mais curtos, apresentam apêndices e ramificações.

Morphoanatomy of the stem in Cyperaceae

Cyperaceae are usually perennial, with underground stems mainly rhizomatous, however, other stem types may also occur, such as corms and tubers. The underground stems of five Cyperaceae species were examined. Cyperus rotundus and Fuirena umbellata have plagiotropic rhizomes, while C. esculentus, C. odoratus, Hypolytrum schraderianum and Bulbostylis paradoxa have orthotropic rhizomes. Corms occur in C. rotundus and C. esculentus, and stolons in C. esculentus. The primary body originates from the activity of the apical meristem and later, from the primary thickening meristem (PTM). Secondary growth results from secondary thickening meristem (STM) activity, and occurs in rhizomes of H. schraderianum, B. paradoxa, C. odotarus and F. umbellata. The procambium and the PTM give rise to collateral bundles in H. schraderianum, and amphivasal bundles in the remaining species. The STM gives rise to the vascular system with the associated phloem and xylem. According to our results, the concept of stem type in Cyperaceae depends on external morphology, function, life phase, activity of the thickening meristems and the relative amount of parenchyma.

Morphological architecture of Actinocephalus (Koern.) sano (Eriocaulaceae-Poales)

Actinocephalus exhibits perhaps more diversity in habit than any other genus of Eriocaulaceae. This variation is largely a result of differences in the arrangement of the paraclades. Based on the analysis of stem architecture of all 25 species of Actinocephalus, the following patterns were established: (1) leaf rosette, with no elongated axis, instead the axillary paraclades originating directly from the short aerial stem, (2) rosette axis continuing into an elongated axis with spirally arranged paraclades, (3) an elongated axis originating from a rhizome, with ramified paraclades, and (4) an elongated axis originating from a short aerial stem, with paraclades arranged in a subwhorl. The elongated axis exhibits indeterminate growth only in pattern 4. Patterns 3 and 4 are found exclusively in Actinocephalus; pattern I occurs in many other genera of Eriocaulaceae, while pattern 2 is also found in Syngonanthus and Paepalanthus. Anatomically, each stem structure (i.e., paraclade, elongated axis, short aerial stem, rhizome) is thickened in a distinctive way and this can be used to distinguish them. Specifically, elongated axes and paraclades lack thickening, thickening of short aerial stems results from the primary thickening meristem and/or the secondary thickening meristem. Thickening of rhizomes results from the activity of the primary thickening meristem. (c) 2008 Elsevier GmbH. All rights reserved.

Morphological architecture of Actinocephalus (Koern.) sano (Eriocaulaceae-Poales)

Actinocephalus exhibits perhaps more diversity in habit than any other genus of Eriocaulaceae. This variation is largely a result of differences in the arrangement of the paraclades. Based on the analysis of stem architecture of all 25 species of Actinocephalus, the following patterns were established: (1) leaf rosette, with no elongated axis, instead the axillary paraclades originating directly from the short aerial stem, (2) rosette axis continuing into an elongated axis with spirally arranged paraclades, (3) an elongated axis originating from a rhizome, with ramified paraclades, and (4) an elongated axis originating from a short aerial stem, with paraclades arranged in a subwhorl. The elongated axis exhibits indeterminate growth only in pattern 4. Patterns 3 and 4 are found exclusively in Actinocephalus; pattern I occurs in many other genera of Eriocaulaceae, while pattern 2 is also found in Syngonanthus and Paepalanthus. Anatomically, each stem structure (i.e., paraclade, elongated axis, short aerial stem, rhizome) is thickened in a distinctive way and this can be used to distinguish them. Specifically, elongated axes and paraclades lack thickening, thickening of short aerial stems results from the primary thickening meristem and/or the secondary thickening meristem. Thickening of rhizomes results from the activity of the primary thickening meristem. (c) 2008 Elsevier GmbH. All rights reserved.

Morfoanatomia dos órgãos vegetativos de Smilax polyantha Griseb. (Smilacaceae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Secondary growth of the Arabidopsis hypocotyl-vascular development in dimensions.

The secondary thickening of plant organs in extant dicotyledons is a massive growth process that constitutes the major carbon sink in perennial, woody plants. Yet, our understanding of its molecular genetic control has been mostly obtained by its analysis in an herbaceous annual model, Arabidopsis. Recent years have seen increased interest in this somewhat under-researched topic, and various (non-)cell autonomous factors that guide the extent and vascular patterning of secondary growth have been identified. Concomitantly, a more detailed understanding of vascular differentiation processes has been obtained through analyses of primary growth, mostly in the root meristem. A future challenge will be the integration of these patterning and differentiation modules together with cambial activity into the 4-dimensional frame of secondary thickening.

Meristematic activity of the Endodermis and the Pericycle in the primary thickening in monocotyledons: considerations on the PTM

A proposta deste trabalho é mostrar uma nova interpretação do meristema de espessamento primário em monocotiledôneas. Anatomia dos órgãos vegetativos das seguintes espécies foi examinada: Cephalostemon riedelianus (Rapataceae), Cyperus papyrus (Cyperaceae), Lagenocarpus rigidus, L. Junciformis (Cyperaceae), Echinodorus paniculatus (Alismataceae) and Zingiberofficinale (Zingiberaceae). A atividade meristemática da endoderme foi observada nas raizes de todas as espécies, no caule de Cyperus, Cephalostemum e Lagenocarpus rigidus, e no traço foliar de Cyperus e folha de Echinodorus. Considerando a continuidade dos tecidos através da raiz, caule e folha, as autoras concluem que no caule o periciclo permanece ativo durante a vida da planta, como um gerador de tecidos vasculares. O Meristema de Espessamento Primário é o periciclo em fase meristemática, juntamente com a endoderme e suas derivadas (ou apenas o periciclo). Próximo ao ápice caulinar, esses tecidos se assemelham a um único meristema, dando origem ao córtex interno e aos tecidos vasculares.

The meristematic activity of the endodermis and the pericycle and its role in the primary thickening of stems in monocotyledonous plants

Background: It had long been thought that a lateral meristem, the so-called primary thickening meristem (PTM) was responsible for stem thickening in monocotyledons. Recent work has shown that primary thickening in the stems of monocotyledons is due to the meristematic activity of both the endodermis and the pericycle. Aims: The aim of this work is to answer a set of questions about the developmental anatomy of monocotyledonous plants: (1) Do the stem apices of monocots have a special meristematic tissue, the PTM? (2) Are the primary tissues of the stem the same as those of the root? (3) Is there good evidence for the formation of both the cortex and the vascular tissue from a single meristem, the PTM, in the shoot and from two distinguishable meristems in the root? (4) If the PTM forms only the cortex, what kind of meristem forms the vascular tissue? Methods: Light microscopy was used to examine stem and root anatomy in 16 species from 10 monocotyledonous families. Results: It was observed that radially aligned cortical cells extend outwards from endodermal initial cells in the cortex of the roots and the stems in all the species. The radial gradation in size observed indicates that the cortical cells are derivatives of a meristematic endodermis. In addition, perfect continuity was observed between the endodermis of the root and that of the stem. Meristematic activity in the pericycle gives rise to cauline vascular bundles composed of metaxylem and metaphloem. Conclusion: No evidence was obtained for the existence in monocotyledons of a PTM. Monocotyledons appear to resemble other vascular plants in this respect.

Morpho-Anatomical Analysis of the Rhizome in Species of Scleria Berg. (Cyperaceae) from Serra do Cipó (MG)

Aspects related to the nature of stem thickening in monocotyledons have been the subject of many studies. Primary thickening has been attributed to the Primary Thickening Meristem (PTM). According to most authors, it gives rise, besides the adventitious roots, to the vascular tissues and part of the cortex. In other words, it has centripetal and centrifugal activity. For some authors, however, it gives rise only to the vascular system, and for others, only to part of the cortex. However, this work demonstrated that PTM corresponds to the pericycle in the meristematic phase or to the pericycle associated with the endodermis, also with meristematic activity. It was observed that the pericycle was responsible for the formation of the vascular system of the rhizome and of the adventitious roots; the endodermis gave rise to cell layers with radial disposition which comprised the inner portion of the stem cortex, and which corresponded to the region known as the derivatives of the meristematic endodermis (DME). A continuity was also demonstrated between the tissues of the stem and root in species of Scleria Berg. (Cyperaceae).

Phylogeny and evolution of anomalous roots in Daviesia (Fabaceae : Mirbelieae)

The phylogeny of the Australian legume genus Daviesia was estimated using sequences of the internal transcribed spacers of nuclear ribosomal DNA. Partial congruence was found with previous analyses using morphology, including strong support for monophyly of the genus and for a sister group relationship between the clade D. pachyloma and the rest of the genus. A previously unplaced bird-pollinated species, anceps + D. D. epiphyllum, was well supported as sister to the only other bird-pollinated species in the genus, D. speciosa, indicating a single origin of bird pollination in their common ancestor. Other morphological groups within Daviesia were not supported and require reassessment. A strong and previously unreported sister clade of Daviesia consists of the two monotypic genera Erichsenia and Viminaria. These share phyllode-like leaves and indehiscent fruits. The evolutionary history of cord roots, which have anomalous secondary thickening, was explored using parsimony. Cord roots are limited to three separate clades but have a complex history involving a small number of gains (most likely 0-3) and losses (0-5). The anomalous structure of cord roots ( adventitious vascular strands embedded in a parenchymatous matrix) may facilitate nutrient storage, and the roots may be contractile. Both functions may be related to a postfire resprouting adaptation. Alternatively, cord roots may be an adaptation to the low-nutrient lateritic soils of Western Australia. However, tests for association between root type, soil type, and growth habit were equivocal, depending on whether the variables were treated as phylogenetically dependent (insignificant) or independent ( significant).

Use of anatomical root markers for species identification in Catasetum (Orchidaceae) at the Portal da Amazônia region, MT, Brazil

Orchidaceae is one of the largest botanical families, with approximately 780 genera. Among the genera of this family, Catasetum currently comprises 166 species. The aim of this study was to characterize the root anatomy of eight Catasetum species, verifying adaptations related to epiphytic habit and looking for features that could contribute to the vegetative identification of such species. The species studied were collected at the Portal da Amazônia region, Mato Grosso state, Brazil. The roots were fixed in FAA 50, cut freehand, and stained with astra blue/fuchsin. Illustrations were obtained with a digital camera mounted on a photomicroscope. The roots of examined species shared most of the anatomical characteristics observed in other species of the Catasetum genus, and many of them have adaptations to the epiphytic habit, such as presence of secondary thickening in the velamen cell walls, exodermis, cortex, and medulla. Some specific features were recognized as having taxonomic application, such as composition of the thickening of velamen cell walls, ornamentation of absorbent root-hair walls, presence of tilosomes, composition and thickening of the cortical cell walls, presence of mycorrhizae, endodermal cell wall thickening, the number of protoxylem poles, and composition and thickening of the central area of the vascular cylinder. These traits are important anatomical markers to separate the species within the genus and to generate a dichotomous identification key for Catasetum. Thus, providing a useful tool for taxonomists of this group

Morphology and anatomy of the vegetative organs and scapes from Aphorocaulon (Paepalanthus, Eriocaulaceae)

Estudou-se a anatomia de raízes, caules, folhas e escapos de espécies de Paepalanthus subseção Aphorocaulon. Estas plantas apresentam caules reduzidos com folhas em roseta, de onde crescem os paracládios (sistemas de inflorescências). As espécies apresentam raízes com epiderme unisseriada e córtex com células isodiamétricas. Tanto os caules reduzidos como os paracládios apresentam espessamento resultante da atividade do periciclo, denominado Meristema de Espessamento Primário (MEP). Ambos apresentam estrutura anatômica semelhante. Os escapos apresentam endoderme descontínua, periciclo sinuoso, o córtex apresenta costelas salientes (5-6). As folhas apresentam células epidérmicas alongadas no sentido longitudinal com paredes levemente espessadas, estômatos somente na face abaxial, com câmara subestomática especializada, feixes vasculares colaterais com bainha dupla. Essas estruturas anatômicas são comuns para as espécies da subseção Aphorocaulon. Algumas características anatômicas observadas nestas espécies são típicas de plantas que crescem nos campos rupestres.

Analysis of secondary growth in the Arabidopsis shoot reveals a positive role of jasmonate signalling in cambium formation.

After primary growth, most dicotyledonous plants undergo secondary growth. Secondary growth involves an increase in the diameter of shoots and roots through formation of secondary vascular tissue. A hallmark of secondary growth initiation in shoots of dicotyledonous plants is the initiation of meristematic activity between primary vascular bundles, i.e. in the interfascicular regions. This results in establishment of a cylindrical meristem, namely the vascular cambium. Surprisingly, despite its major implications for plant growth and the accumulation of biomass, the molecular regulation of secondary growth is only poorly understood. Here, we combine histological, molecular and genetic approaches to characterize interfascicular cambium initiation in the Arabidopsis thaliana inflorescence shoot. Using genome-wide transcriptional profiling, we show that stress-related and touch-inducible genes are up-regulated in stem regions where secondary growth takes place. Furthermore, we show that the products of COI1, MYC2, JAZ7 and the touch-inducible gene JAZ10, which are components of the JA signalling pathway, are cambium regulators. The positive effect of JA application on cambium activity confirmed a stimulatory role of JA in secondary growth, and suggests that JA signalling triggers cell divisions in this particular context.