Pericarp ontogeny and histochemistry of the exotesta and pseudocaruncle of <i>Euphorbia milii</i> (Euphorbiaceae)

Demarco, Diego; Carmello-Guerreiro, Sandra Maria

doi:10.1590/2175-7860201162304

Abstract

Several types of fruit occur in Euphorbiaceae, notably the explosively dehiscent dry fruit, and different seed-coat anatomies with taxonomic importance. This paper aims to describe the pericarp ontogeny and structure in Euphorbia milii Desmoul., and evaluate the presence of the secretory exotesta and caruncle. The fruit is a schizocarp, whose the pericarp development begins with a periclinal division of the inner epidermal cells. The derived cells divide, forming about four layers of obliquely elongated cells. Then, the adjacent parenchyma cells elongate, giving rise to a palisade layer and finally, the cells between this layer and the vascular strands undergo mitosis, originating about four layers of elongated cells perpendicularly to the inner oblique cells. These three zones lignify, while the region between the vascular strands and the exocarp, where idioblasts, hypodermis and laticifers are present do not show significant changes. Before the dehiscence, a lysis of cells of the septa and the desiccation of the fruit occur, which causes contraction of the non-lignified tissues and tension between the lignified zones, promoting rupture of each mericarp from central columella and on the dorsal strand, ejecting the seeds. The seeds have pseudocaruncle and the exotesta secretes mucilage, facilitating their imbibition.

Key words:
anatomy; crown-of-thorns; development; fruit; seed

Resumo

Diversos tipos de frutos ocorrem em Euphorbiaceae, destacando-se os secos de deiscência explosiva e sementes com envoltórios de diferentes estruturas e importância taxonômica. O objetivo deste trabalho foi descrever a ontogênese do pericarpo de Euphorbia milii Desmoul. e avaliar, em termos histoquímicos, a exotesta secretora e a presença de carúncula. O fruto é um esquizocarpo, cujo desenvolvimento do pericarpo se inicia pela divisão periclinal das células da epiderme interna. As derivadas dividem-se, formando cerca de quatro camadas de células alongadas obliquamente. Em seguida, as células parenquimáticas adjacentes alongam-se, formando uma camada em paliçada e então, as células entre a camada em paliçada e os feixes vasculares dividem-se, originando cerca de quatro camadas de células alongadas perpendicularmente às células oblíquas internas. Estas três zonas lignificam-se, enquanto a região entre os feixes vasculares e o exocarpo, que possui idioblastos, hipoderme e laticíferos, não apresenta mudanças significativas. Antes da deiscência ocorre lise das células dos septos e dessecamento do fruto, o qual gera contração dos tecidos não lignificados e tensão entre as zonas lignificadas, rompendo os mericarpos em relação à columela central e na região do feixe dorsal, lançando as sementes. As sementes possuem pseudocarúncula e a exotesta secreta mucilagem, facilitando sua embebição.

Palavras-chave:
anatomia; coroa-de-cristo; desenvolvimento; fruto; semente

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References

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Boesewinkel, F.D. ; & Bouman, F. 1984. The seed: structure. In: Johri, B.M. Embryology of angiosperms. Springer, Berlin. Pp. 567-610.
Bor, J. & Bouman, F. 1974. Development of ovule and integuments in Euphorbia milii and Codiaeum variegatum Phytomorphology 24: 280-296.
Cain, A.J. 1947. The use of Nile Blue in the examination of lipids. Quarterly Journal of Microscopical Science 88: 383-392.
Carlquist, S. 1966. The biota of long-distance dispersal. III. Loss of dispersibility in the Hawaiian Flora. Brittonia 18: 310-335.
Esser, H. 2003. Fruit characters in Malesian Euphorbiaceae. Telopea 10: 169-177.
Fahn, A. 1990. Plant anatomy. 4ed. Pergamon Press, Oxford. 588p.
Gabe, M. 1968. Techniques histologiques. Masson & Cie, Paris. 1113p.
Ganter, P. & Jollés, G. 1969, 1970. Histologie normale et pathologique. 2 vol. Gauthier-Villars, Paris. 1904p.
Gerlach, D. 1984. Botanische Mikrotechnik: eine Einführung. 3ed. Georg Thieme, Stuttgart. 311p.
Gerrits, P.O. 1991. The application of glycol methacrylate in histotechnology; some fundamental principles. Department of Anatomy and Embryology, State University Groningen, Groningen. 80p.
Gregory, M.; & Baas, P. 1989. A survey of mucilage cells in vegetative organs of the dicotyledons. Israel Journal of Botany 38: 125-174.
High, O.B. 1984. Lipid histochemistry. Oxford University Press, New York. 68p.
Hyde, B.B. 1970. Mucilage-producing cells in the seed coat of Plantago ovata: developmental fine structure. American Journal of Botany 57: 1197-1206.
Jimenez, M.S.; & Morales, D. 1978. Histología floral y desarrollo del fruto de Euphorbia atropurpurea Brouss. Vieraea 8: 131-143.
Johansen, D.A. 1940. Plant microtechnique. McGraw-Hill, New York. 523p.
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Lillie, R.D. 1965. Histopathologic technic and practical histochemistry. 3ed. McGraw-Hill, New York. 715p.
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Muzik, T.J. 1954. Development of fruit, seed, embryo, and seedling of Hevea brasiliensis American Journal of Botany 41:39-43.
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Pearse, A.G.E. 1985. Histochemistry: theoretical and applied. 4ed. Vol. 2. C. Livingstone, Edinburgh. 624p.
Pizzolato, T.D. 1977. Staining of Tilia mucilages with Mayer's tannic acid- ferric chloride. Bulletin of the Torrey Botanical Club 104: 277-279.
Sablon, L. du 1884. Recherches sur la déhiscence des fruits a péricarpe sec. Annales des Sciences Naturelles. Botanique et Biologie Vegetale 18: 5-104.
Simon, J.; Molero, J. & Blanché, C. 1992. Fruit and seed morphology of Taxonomic implications. Collectanea Botanica 21: 211-242.
Singh, R.P. 1969. Structure and development of seeds in Euphorbia helioscopia Botanical Magazine Tokyo 82: 287-293.
Subils, R. 1977. Las especies de Euphorbia de la Republica Argentina. Kurtziana 10: 83-248.
Tokuoka, T.; & Tobe, H. 2002. Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications. Journal of Plant Research 115: 361-374.
Toledo, A.P. 1963. Anatomia e desenvolvimento ontogenético do fruto e da semente de mandioca. Bragantia 22: 71-76.
Webster, G.L. 1994. Classification of the Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 3-32.
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Publication Dates

Publication in this collection
Jul-Sep 2011

History

Received
01 Dec 2010
Accepted
23 May 2011

This is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

[1] Berg, R.Y. 1975. Myrmecochorous plants in Australia and their dispersal by ants. Australian Journal of Botany 23 : 475-508.

[2] Boesewinkel, F.D. ; & Bouman, F. 1984. The seed: structure. In: Johri, B.M. Embryology of angiosperms. Springer, Berlin. Pp. 567-610.

[3] Bor, J. & Bouman, F. 1974. Development of ovule and integuments in Euphorbia milii and Codiaeum variegatum Phytomorphology 24: 280-296.

[4] Cain, A.J. 1947. The use of Nile Blue in the examination of lipids. Quarterly Journal of Microscopical Science 88: 383-392.

[5] Carlquist, S. 1966. The biota of long-distance dispersal. III. Loss of dispersibility in the Hawaiian Flora. Brittonia 18: 310-335.

[6] Esser, H. 2003. Fruit characters in Malesian Euphorbiaceae. Telopea 10: 169-177.

[7] Fahn, A. 1990. Plant anatomy. 4ed. Pergamon Press, Oxford. 588p.

[8] Gabe, M. 1968. Techniques histologiques. Masson & Cie, Paris. 1113p.

[9] Ganter, P. & Jollés, G. 1969, 1970. Histologie normale et pathologique. 2 vol. Gauthier-Villars, Paris. 1904p.

[10] Gerlach, D. 1984. Botanische Mikrotechnik: eine Einführung. 3ed. Georg Thieme, Stuttgart. 311p.

[11] Gerrits, P.O. 1991. The application of glycol methacrylate in histotechnology; some fundamental principles. Department of Anatomy and Embryology, State University Groningen, Groningen. 80p.

[12] Gregory, M.; & Baas, P. 1989. A survey of mucilage cells in vegetative organs of the dicotyledons. Israel Journal of Botany 38: 125-174.

[13] High, O.B. 1984. Lipid histochemistry. Oxford University Press, New York. 68p.

[14] Hyde, B.B. 1970. Mucilage-producing cells in the seed coat of Plantago ovata: developmental fine structure. American Journal of Botany 57: 1197-1206.

[15] Jimenez, M.S.; & Morales, D. 1978. Histología floral y desarrollo del fruto de Euphorbia atropurpurea Brouss. Vieraea 8: 131-143.

[16] Johansen, D.A. 1940. Plant microtechnique. McGraw-Hill, New York. 523p.

[17] Jordan, M.S. & Hayden, W.J. 1992. A survey of mucilaginous testa in Chamaesyce Collectanea Botanica 21: 79-89.

[18] Jordan, L.S.; Jordan, J.L. & Jordan, C.M. 1985. Changes induced by water on Euphorbia supina seed coat structures. American Journal of Botany 72: 1530-1536.

[19] Kapil, R.N.; Bor, J.; & Bouman, F. 1980. Seed appendages in angiosperms. I. Introduction. Botanische Jahrbücher für Systematik 101: 555-573.

[20] Lavialle, P. & Delacroix, J. 1922a. La paroi du pistil et du fruit dans le genre Euphorbia Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences 175: 179-181.

[21] Lavialle, P. & Delacroix, J. 1922b. Caractères de l'endocarpe dans le genre Euphorbia Bulletin de la Société Botanique de France 69: 523-527.

[22] Lavialle, P. & Delacroix, J. 1922c. Caractères histologiques du péricarpe et déhiscence du fruit chez les Euphorbes. Bulletin de la Société Botanique de France 69: 585-590.

[23] Lillie, R.D. 1965. Histopathologic technic and practical histochemistry. 3ed. McGraw-Hill, New York. 715p.

[24] Lisci, M.; Bianchini, M. & Pacini, E. 1996. Structure and function of the elaiosome in some angiosperm species. Flora 191: 131-141.

[25] Lisci, M. & Pacini, E. 1997. Fruit and seed structural characteristics and seed dispersal in Mercurialis annua L. (Euphorbiaceae). Acta Societatis Botanicorum Poloniae 66: 379-386.

[26] Liszt, K.; Sárkány, S.; Kadej, J.; & Kovács, A. 1981. Light and electron microscopic observations in connection with the developing pistil and seed-appendix (caruncle) of Ricinus communis L. Acta Societatis Botanicorum Poloniae 50: 345-348.

[27] McManus, J.F.A. 1948. Histological and histochemical uses of periodic acid. Stain Technology 23: 99-108.

[28] Muzik, T.J. 1954. Development of fruit, seed, embryo, and seedling of Hevea brasiliensis American Journal of Botany 41:39-43.

[29] Narbona, E.; Arista, M.; & Ortiz, P.L. 2005. Explosive seed dispersal in two perennial Mediterranean Euphorbia species (Euphorbiaceae). American Journal of Botany 92: 510-516.

[30] O'Brien, T.P.; Feder, N.; & McCully, M.E. 1964. Polychromatic staining of plant cell walls by toluidine blue O. Protoplasma 59: 368-373.

[31] Oliveira, J.H.G.; & Oliveira, D.M.T. 2009. Morfoanatomia e ontogênese do pericarpo de Manihot caerulescens Pohl e M. tripartita Müll.Arg. (Euphorbiaceae). Revista Brasileira de Botânica 32: 117-129.

[32] Pearse, A.G.E. 1985. Histochemistry: theoretical and applied. 4ed. Vol. 2. C. Livingstone, Edinburgh. 624p.

[33] Pizzolato, T.D. 1977. Staining of Tilia mucilages with Mayer's tannic acid- ferric chloride. Bulletin of the Torrey Botanical Club 104: 277-279.

[34] Sablon, L. du 1884. Recherches sur la déhiscence des fruits a péricarpe sec. Annales des Sciences Naturelles. Botanique et Biologie Vegetale 18: 5-104.

[35] Simon, J.; Molero, J. & Blanché, C. 1992. Fruit and seed morphology of Taxonomic implications. Collectanea Botanica 21: 211-242.

[36] Singh, R.P. 1969. Structure and development of seeds in Euphorbia helioscopia Botanical Magazine Tokyo 82: 287-293.

[37] Subils, R. 1977. Las especies de Euphorbia de la Republica Argentina. Kurtziana 10: 83-248.

[38] Tokuoka, T.; & Tobe, H. 2002. Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications. Journal of Plant Research 115: 361-374.

[39] Toledo, A.P. 1963. Anatomia e desenvolvimento ontogenético do fruto e da semente de mandioca. Bragantia 22: 71-76.

[40] Webster, G.L. 1994. Classification of the Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 3-32.

[41] Werker, E. 1997. Seed anatomy (Handbuch der Pflanzenanatomie; Bd. 10, Teil 3). Gebrüder Borntraeger, Berlin. 424p.

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