The origin of new roots from cut seeds of <i>Eugenia</i> species

Studies on seed germination in Eugenia species after embryo fractionation have demonstrated that cotyledon cells have capacity for de-differentiation and consequent production of roots. However, there is no information about the origin of those new roots. Thus, the aim of this study was to characterize anatomically the cotyledon regions of seeds of five species of Eugenia to elucidate the tissue that originates such roots. Seeds were sectioned across the hillum region and immediately placed to germinate. As soon as the fractions of these cut seeds develop roots they were fixed and processed to the usual techniques for light microscopy. The adventitious roots originated only when the seed was fractioned, never occurring spontaneously in uninjured seeds. Adventitious roots were formed from perivascular parenchyma cells, located close to the injured region of the cotyledons. These cells divided periclinally and proliferated, giving rise to a root meristem. Therefore, new seedlings probably have a different genome than the mother plant, but they would be identical to the embryo that was a result of fertilization.

Index terms:
cell de-differentiation; fractionation; perivascular cells; plant anatomy

Authorship

Liliana Ferreira Delgado

Secretaria Municipal de Educação de São Paulo, EMEF Primo Pascoli Melare, Prof., 02882-160 - São Paulo, SP, Brasil.Secretaria Municipal de Educação de São PauloBrazilSão Paulo, SP, BrazilSecretaria Municipal de Educação de São Paulo, EMEF Primo Pascoli Melare, Prof., 02882-160 - São Paulo, SP, Brasil.

http://orcid.org/0000-0002-1241-3887

Simone Pádua Teixeira

Universidade de São Paulo, Faculdade de Ciências Farmacêuticas de Ribeirão Preto, 14040-903 - Ribeirão Preto, SP, Brasil.Universidade de São PauloBrazilRibeirão Preto, SP, BrazilUniversidade de São Paulo, Faculdade de Ciências Farmacêuticas de Ribeirão Preto, 14040-903 - Ribeirão Preto, SP, Brasil.

http://orcid.org/0000-0003-4993-101X

Sandra Maria Carmello-Guerreiro

Universidade Estadual de Campinas, Instituto de Biologia, Departamento de Biologia Vegetal, 13083-862 - Campinas, SP - Brasil.Universidade Estadual de CampinasBrazilCampinas, SP, BrazilUniversidade Estadual de Campinas, Instituto de Biologia, Departamento de Biologia Vegetal, 13083-862 - Campinas, SP - Brasil.

http://orcid.org/0000-0002-0348-1511

Claudio José Barbedo ^**Corresponding author E-mail: cjbarbedo@yahoo.com.br

Instituto de Pesquisas Ambientais, Núcleo de Conservação da Biodiversidade, 04301-012 - São Paulo, SP, Brasil.Instituto de Pesquisas AmbientaisBrasilSão Paulo, SP, BrasilInstituto de Pesquisas Ambientais, Núcleo de Conservação da Biodiversidade, 04301-012 - São Paulo, SP, Brasil.

http://orcid.org/0000-0002-4417-7495

*Corresponding author
E-mail: cjbarbedo@yahoo.com.br

SCIMAGO INSTITUTIONS RANKINGS

Figures | Tables

Figures (4)
Tables (1)

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Figure 1
Fruits, seeds, embryos and aspects of seed fraction germination of Eugenia species. A-D - Eugenia brasiliensis, A - Fruit, B - Seed, C- Embryo, D - germinated seed fraction; E-H - E. involucrata, E - Fruit, F - Seed, G - Embryo, H - germinated seed fraction; I-L - E. cerasiflora, I - Fruit, J - Seed, K- Embryo, L - germinated seed fraction; M-P - E. pyriformis, M - Fruit, N - Seed, O - Embryo, P - germinated seed fraction; Q-T - E. uniflora, Q - Fruit, R - Seed, S - Embryo, T - germinated seed fraction.

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Figure 2
Whole seeds of Eugenia cerasiflora (A) and E. involucrata (B) with a small protrusion (arrow) surrounded by a circular depression which allows to distinguish the hypocotyl-radicle axis by naked eye, not so evident in E. uniflora (C). D: Vascular bundles (arrow) in the peripheral area of cotyledons showed by a transversal section of E. involucrata embryo from cut seed. E: Multiple adventitious roots from the cut embryo of E. uniflora from cut seed. F: Schematic illustration of transversal (CT) and longitudinal (CL) sections of E. involucrata seed showing the vascular bundles (FV) and hillum (H). Anatomical transversal sections of the hypocotyl-radicle axis (arrows) of seeds from immature fruits of E. cerasiflora (G), E. involucrata (H) and E. uniflora (I). J: Transversal section of cut E. uniflora seed after four days from the beginning of germination (the arrow shows the greater celular proliferation of the vascular bundle than the other marked by an asterisk).

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Figure 3
Anatomical sections of cotyledons of Eugenia embryos. A: Two cotyledons of E. uniflora embryo identified by their uniseriate epidermis; B: Uniseriate epidermis and union of cotyledons of E. involucrata embryo; C: Detail of a secretory cavity with the secretory multiseriate epithelium in E. uniflora. Note that the cotyledon epidermis is uniseriate; D: Uniseriate epidermis with cuticle in E. cerasiflora embryo; E: Wound-healing tissue after larval predation in E. cerasiflora embryo; F: Wound-healing tissue after cut in E. cerasiflora embryo; G: Starch in the cotyledon of E. cerasiflora; H: Rudimentary embryonic axis in seed from mature fruit of E. uniflora.

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Figure 4
Anatomical sections of cotyledons of Eugenia embryos. A-E: Adventitious root from the perivascular parenchyma cells after cutting the embryo of Eugenia involucrata (A), E. uniflora (B), E. brasiliensis (C), E. cerasiflora (D) and E. pyriformis (E). F-H: Periclinal divisions and proliferation of perivascular cells in embryos of E. cerasiflora (F), E. uniflora (G), E. involucrata (H) after nine days of cutting seeds. I: Adventitious root in E. uniflora embryo after seed cut.

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Table 1
Regenerative ability described for seeds of Eugenia species.

Figure 1 Fruits, seeds, embryos and aspects of seed fraction germination of Eugenia species. A-D - Eugenia brasiliensis, A - Fruit, B - Seed, C- Embryo, D - germinated seed fraction; E-H - E. involucrata, E - Fruit, F - Seed, G - Embryo, H - germinated seed fraction; I-L - E. cerasiflora, I - Fruit, J - Seed, K- Embryo, L - germinated seed fraction; M-P - E. pyriformis, M - Fruit, N - Seed, O - Embryo, P - germinated seed fraction; Q-T - E. uniflora, Q - Fruit, R - Seed, S - Embryo, T - germinated seed fraction.

Figure 2 Whole seeds of Eugenia cerasiflora (A) and E. involucrata (B) with a small protrusion (arrow) surrounded by a circular depression which allows to distinguish the hypocotyl-radicle axis by naked eye, not so evident in E. uniflora (C). D: Vascular bundles (arrow) in the peripheral area of cotyledons showed by a transversal section of E. involucrata embryo from cut seed. E: Multiple adventitious roots from the cut embryo of E. uniflora from cut seed. F: Schematic illustration of transversal (CT) and longitudinal (CL) sections of E. involucrata seed showing the vascular bundles (FV) and hillum (H). Anatomical transversal sections of the hypocotyl-radicle axis (arrows) of seeds from immature fruits of E. cerasiflora (G), E. involucrata (H) and E. uniflora (I). J: Transversal section of cut E. uniflora seed after four days from the beginning of germination (the arrow shows the greater celular proliferation of the vascular bundle than the other marked by an asterisk).

Figure 3 Anatomical sections of cotyledons of Eugenia embryos. A: Two cotyledons of E. uniflora embryo identified by their uniseriate epidermis; B: Uniseriate epidermis and union of cotyledons of E. involucrata embryo; C: Detail of a secretory cavity with the secretory multiseriate epithelium in E. uniflora. Note that the cotyledon epidermis is uniseriate; D: Uniseriate epidermis with cuticle in E. cerasiflora embryo; E: Wound-healing tissue after larval predation in E. cerasiflora embryo; F: Wound-healing tissue after cut in E. cerasiflora embryo; G: Starch in the cotyledon of E. cerasiflora; H: Rudimentary embryonic axis in seed from mature fruit of E. uniflora.

Figure 4 Anatomical sections of cotyledons of Eugenia embryos. A-E: Adventitious root from the perivascular parenchyma cells after cutting the embryo of Eugenia involucrata (A), E. uniflora (B), E. brasiliensis (C), E. cerasiflora (D) and E. pyriformis (E). F-H: Periclinal divisions and proliferation of perivascular cells in embryos of E. cerasiflora (F), E. uniflora (G), E. involucrata (H) after nine days of cutting seeds. I: Adventitious root in E. uniflora embryo after seed cut.

Table 1 Regenerative ability described for seeds of Eugenia species.

Species	Information	Source
E. pyriformis	Seeds cut to leave 1/4 of their original size can germinate and produce normal seedlings	1
	The embryo fills the entire space delimited by the integument and the axis is less than 1.0 mm long; the fleshy cotyledons vary from 1.0 to 2.0 cm	2
	Fragmented seeds show potential for root and seedling regeneration, but germination inhibits the formation of new roots and shoots	3
	Seeds can germinate and produce normal seedlings after the removal of 3/4 of their mass, even for immature and germinating ones	4
	The capacity to produce new seedlings reduces with the reduction in seed size	5
	Germination and normal seedlings are greater for entire seeds than the cut ones	6
	Seeds have high capacity to producing normal seedlings even from fruits at the beginning of development and after some seed reserve material is lost	7
E. involucrata	A batch in which seeds are cut into two parts can germinate more than 100%, but do not produce more than 100% of normal seedlings	8
	Seeds can germinate and produce normal seedlings after the removal of 3/4 of their mass, even for immature and germinating ones	4
	The uncut seeds have higher germination percentage than the cut ones	9
	Seeds have high capacity to producing normal seedlings even from fruits at the beginning of development and after some seed reserve material is lost	7
	Seeds are able to produce new roots and seedlings until the third radicle/seedling produced is removed	10
	Roots and seedlings continued to develop from apparently water stress necrotic tissue on the seed surface	11
E. brasiliensis	A batch in which seeds are cut into two parts can germinate more than 100%, but do not produce more than 100% of normal seedlings	8
	Seeds can germinate and produce normal seedlings after the removal of 3/4 of their mass, even for immature and germinating ones	4
	Seed germination inhibits the formation of new roots or seedlings in the seed	12
	Seeds have high capacity to producing normal seedlings even from fruits at the begining of development and after some seed reserve material is lost	7
	Seeds are able to produce new roots and seedlings until the third radicle/seedling produced is removed	10
E. uniflora	A batch in which seeds are cut into two parts can germinate more than 100%, but do not produce more than 100% of normal seedlings	8
	Seeds can germinate and produce normal seedlings after the removal of 3/4 of their mass, even for immature and germinating ones	4
	Seed germination starts the inhibition of the formation of new roots or seedlings in the seed and seed incision can block the action of these inhibitors	12
	Seeds have high capacity to producing normal seedlings even from fruits at the beginning of development and after some seed reserve material is lost	7
E. stipitata	Seeds show a high capacity for regeneration; even when cut in the middle or in the meristematic zone, they still form seedlings	13
	Cut seeds cut at the meristematic protuberance make normal seedlings, with the same characteristics of intact seed germination	14
	When seedlings are detached from the seed, the ‘resown’ seeds produce a second, normal seedling within 9 months	15
E. cerasiflora	A batch in which seeds are cut into four parts can germinate more than 200%	16
	Seeds can germinate and produce normal seedlings after the removal of 3/4 of their mass, even for immature and germinating ones	4
	Seeds have high capacity to producing normal seedlings even from fruits at the beginning of development and after some seed reserve material is lost	7
E. candolleana	It is possible to obtain more than one normal seedling from each seed; after root and shoot removal, regeneration of new roots and seedlings occurred	17
E. umbelliflora	The seeds have high capacity of resume germinability after being cut	16
E. pruinosa	The seeds have high capacity of resume germinability after being cut	16

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The origin of new roots from cut seeds of <i>Eugenia</i> species

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[1] *Corresponding author
E-mail: cjbarbedo@yahoo.com.br