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.O crescimento em espessura do caule de monocotiledônea pode ser primário, ou primário e secundário. A maioria dos autores consideram o espessamento resultante do MEP (Meristema de Espessamento Primário) e do MES (Meristema de Espessamento Secundário). Há divergências de qual seria o meristema responsável pelo espessamento primário. Em Cordyline fruticosa o procâmbio forma feixes vasculares de dois tipos: traços foliares colaterais (com proto e metaxilema e proto e metafloema), e feixes caulinares concêntricos (com metaxilema e metafloema). O procâmbio também forma o periciclo, a camada mais externa do cilindro vascular, constituída por células menores e menos coradas que se dividem irregularmente, formando novos feixes vasculares. O periciclo dá continuidade à atividade procambial, originando somente feixes concêntricos. Concluiu-se ser o periciclo responsável pelo espessamento primário desta espécie. Mais distante do ápice as células pericíclicas passam a sofrer divisões periclinais originando o Meristema de Espessamento Secundário. A análise dos cortes seriados mostra que o periciclo e o MES são contínuos nesta espécie, ficando claro que o periciclo origina oMES. A endoderme é reconhecida, apenas, como a camada mais interna do córtex.Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq

Rhizophores in Rhizophora mangle L: an alternative interpretation of so-called ''aerial roots''

Rhizophora mangle L., one of the most common mangrove species, has an aerial structure system that gives it stability in permanently swampy soils. In fact, these structures, known as "aerial roots" or "stilt roots", have proven to be peculiar branches with positive geotropism, which form a large number of roots when in contact with swampy soils. These organs have a sympodial branching system, wide pith, slightly thickened cortex, collateral vascular bundles, polyarch stele and endarch protoxylem, as in the stem, and a periderm produced by a phellogen at the apex similar to a root cap. They also have the same type of trichosclereid that occurs in the stem, with negative geotropism, unlike true Rhizophora roots, which do not form trichosclereids at all. On the other hand, these branches do not form leaves and in this respect they are similar to roots. These peculiar branches are rhizophores or special root-bearing branches, analogous to those found in Lepidodendrales and other Carboniferous tree ferns that grew in swampy soils

A new interpretation on vascular architecture of the cauline system in Commelinaceae (Commelinales).

The vascular system of monocotyledons, including Commelinaceae, has been studied since the 19th century, but to date, the proposed vascular architecture models consist of schematic representations partially based on the authors' interpretation. One of the greatest difficulties in studying these systems is the large number of vascular bundles and the complexity of their connections, especially in the monocotyledons which have a nodal vascular plexus. In this study, shoot apex samples of 14 species of Commelinaceae were submitted to three-dimensional analyses (confocal microscopy, X-ray microtomography, graphic vectorization, and whole-mount diaphanization), as well as conventional techniques in plant anatomy. Based on the results, a previously unreported category of bundles is described in Commelinaceae for the first time, as well as the fact that peripheral bundles are not interrupted or end blindly in the periphery of the pith, as previously thought. With this new interpretation of the vascular architecture, three patterns of nodal vascular plexus are proposed: 1) in the first pattern the internal nodal vascular plexus (IVP) forms a continuous cylinder and does not connect to the external nodal vascular plexus (EVP); 2) the IVP forms a cylinder divided into two columns and does not connect to the EVP and 3) the IVP forms a cylinder connected to the EVP. The first description of central bundles in the Commelinaceae might suggests their existence in closely related groups, such as the remaining four families of Commelinales (i.e., Haemodoraceae, Hanguanaceae, Philydraceae, and Pontederiaceae), and even in other distantly related groups of monocotyledons

Erratum to: Meristematic activity of the endodermis and the pericycle in the primary thickening in monocotyledons. Considerations on the "PTM" [An Acad Bras Cienc 77(2005): 259-274]

This paper proposes a new interpretation for primary thickening in monocotyledons. The anatomy of the vegetative organs of the following species was examined: Cephalostemon riedelianus (Rapataceae), Cyperus papyrus (Cyperaceae), Lagenocarpus rigidus, L. junciformis (Cyperaceae), Echinodorus paniculatus (Alismataceae) and Zingiber officinale (Zingiberaceae). The endodermis with meristematic activity was observed in the root of all the species, in the stem of Cyperus, Cephalostemum and Lagenocarpus rigidus, and in the leaf trace of Cyperus and leaf of Echinodorus. Considering the continuity of tissues through the root, stem and leaf, the authors conclude that in the stem the pericycle remains active throughout the life of the plant as the generator of the vascular tissue. The "Primary Thickening Meristem" is in fact the pericycle plus the endodermis and its derivatives (or only the pericycle). Close to the stem apex, the assemblage of seems to be a unique meristem, giving rise to the inner cortex and vascular tissues.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.Sao Paulo Research Foundation (FAPESP)Sao Paulo Research Foundation (FAPESP)National Council for Scientific and Technological Development (CNPq)National Council for Scientific and Technological Development (CNPq

Endoderme com atividade meristem\ue1tica em raiz de Canna edulis Kerr-Gawler (Cannaceae)

Canna edulis é uma planta ornamental utilizada em muitos países como fonte alimentar alternativa. O objetivo deste trabalho foi descrever a formação do córtex radicular a partir da análise anatômica da região apical. Na região situada a 220µm do pró-meristema, os tecidos meristemáticos apicais já se apresentam organizados em protoderme, meristema fundamental e procâmbio. Em fase subseqüente na diferenciação celular, a 450µm do pró-meristema, as camadas de células do córtex estão dispostas em fileiras radiais iniciando-se na endoderme. Depois que as iniciais endodérmicas cessam as divisões, adquirem estrias de Caspary. Na raiz, a 1.700µm do pró-meristema, os tecidos primários já se encontram diferenciados, sendo o padrão de distribuição celular observado no córtex de C. edulis característico ao apresentado por outras espécies de Zingiberales. A análise anatômica da região apical levou à constatação que 2/3 do córtex é resultante da atividade meristemática da endoderme e o restante das células corticais são originadas diretamente do meristema fundamental