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Morphology, Anatomy and Germination Response of Heteromorphic Achenes of Anthemis Chrysantha J

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Morphology, Anatomy and Germination Response of Heteromorphic Achenes of Anthemis Chrysantha J Document downloaded from: http://hdl.handle.net/10251/28830 This paper must be cited as: García Breijo, FJ.; Reig Armiñana, J.; Aguado, M.; Martinez-sanchez, JJ.; Franco, JA.; Vicente, MJ. (2011). Morphology, anatomy and germination response of heteromorphic achenes of Anthemis chrysantha J. Gay (Asteraceae), a critically endangered species.. SEED SCIENCE RESEARCH. 21(4):283-294. doi:10.1017/S0960258511000183. The final publication is available at http://dx.doi.org/10.1017/S0960258511000183 Copyright Cambridge University Press Morphology, anatomy and germination response of heteromorphic achenes of Anthemis chrysantha J. Gay (Asteraceae), a critically endangered species Mayra Aguado1, Juan J. Martı´nez-Sa´nchez1*, Jose´ Reig-Armin˜ ana2, Francisco J. Garcı´a-Breijo2,3, Jose´ A. Franco1 and Marı´a J. Vicente1 1Departamento de Produccio´n Vegetal, Instituto de Biotecnologı´a Vegetal, Universidad Polite´cnica de Cartagena, Paseo Alfonso XIII, 48, 30203 Cartagena, Spain; 2Laboratorio ‘Julio Iranzo’ de Anatomı´a Vegetal, Jardı´n Bota´nico de la Universidad de Valencia, C/ Quart, 80, 46008 Valencia, Spain; 3Departamento de Ecosistemas Agroforestales, Universidad Polite´cnica de Valencia, Camino de Vera s/n. 46022, Valencia, Spain Abstract physically impeding germination and hampering imbibition of water. This study demonstrates that Anthemis chrysantha,a ‘Critically Endangered’ annual plant, produces two morphs of achenes: white and dark achenes, which Keywords: Anthemis, endangered species, heteromorphic differ in size, mass, anatomy and germination fruits, pericarp structure, seed dormancy, seed behaviour. Fresh white achenes germinated at all germination temperatures assayed from 10 to 258Cinboth continuous darkness and 12-h photoperiod, ranging between 24% at 258C in darkness and 89% at 12/208C in light, whereas fresh dark achenes did not germinate under any temperature or light conditions. To identify Introduction differences in dormancy type between the two morphs, germination of dry-stored achenes, and achenes The morphology and physiological behaviour of seeds are important features for understanding the pattern of stratified at 5 or 258C for 2 months were tested in seasonal and spatial distributions of species (Silvertown both darkness and light at 5, 15 and 12/208C for dry- and Doust, 1993; Imbert, 2002). Many species of stored and warm-stratified (258C) achenes; and at 15, Asteraceae are known for producing two or more 25 and 12/208C for cold-stratified (58C) achenes. Of different morphs of achenes within a single plant the white achenes, 90% germinated during the cold (Imbert, 2002; Bra¨ndel, 2004), which is generally stratification period. In general, dry storage and warm associated with adaptation to unstable environments stratification did not increase germination compared (Harper, 1977; Venable, 1985). Seed polymorphism can to fresh achenes. However, dark achenes did not affect characteristics such as dispersal capacity, seed germinate under any conditions. Dark achene dor- dormancy, predation, germinability and seedling com- mancy was only broken by mechanical scarification or petition (Harper, 1977; Imbert et al.,1996;Bra¨ndel, 2004). by excising the embryo (germination reached 71%). According to Nikolaeva (1977), in endogenous An anatomical study showed that the mesocarp of dark dormancy some characteristic of the embryo prevents achenes had no intercellular spaces and was much germination, whereas in exogenous dormancy, some thicker and stronger than that of white achenes, characteristic of structures covering the embryo making the entry of water difficult, and also preventing prevents germination. This author defined physical germination by mechanical restriction. This study exogenous dormancy as an impermeability to water of demonstrated that dormancy in the dark achenes is the seed (fruit) coats. Most types of dormancy can be likely caused by the thickness of their pericarp, broken by warm and/or cold stratification, but not physical exogenous dormancy. There have been studies on the morphology of *Correspondence the capitulum and the fruit in order to establish Fax: þ 34 968325433 phylogenetic classifications within the Asteraceae Email: [email protected] (Chehregani and Mahanfar, 2007; Kreitschitz and M. Aguado et al. Valle´s, 2007), and some have described relationships and a denticulate rim or sometimes a short crenulate between fruit morphology and germination capacity auricle (pappus). Under a stereoscopic microscope two (Porras and Mun˜oz, 2000; Imbert, 2002; Bra¨ndel, 2004; principal morphs of achenes are distinguishable Sun et al.,2009),butveryfewhaveaddressed within one fruit head: the rows of the upper section germination in the genus Anthemis (Rashid et al., containing elongated, almost white achenes (white 2007). Anthemis L. is the second largest genus within achenes hereafter), while the achenes of the basal rows the Asteraceae (tribe Anthemidae), with more than 210 are brown–black (dark achenes hereafter) and harder species in the Mediterranean region, south-west Asia than white achenes. Finally, the last basal line is and eastern Africa (Oberprieler, 2001); about 62 species composed of achenes from ligules (only 8–10 per fruit are distributed in Europe and 14 in the Iberian head), which are similar to white achenes but Peninsula. One of these species is A. chrysantha J. Gay, frequently are empty, and were not studied in this an annual plant endemic to North Africa and the work. The proportion of white and dark achenes in a south-east of the Iberian Peninsula. It is only found on fruit head is about 70 and 30%, respectively. The the Algerian coast and, in Europe, only on the coast of percentage of empty white and dark achenes is very Cartagena (Murcia, south-eastern Spain). In Cartagena, variable from one fruit head to another, but the average four populations had been known, but since the in the dark achenes is always lower than in white late 1990s only two have remained, occupying an area achenes (about 10 and 24%, respectively). A represen- of less than 2 ha. A. chrysantha grows in therophytic tative sample of freshly matured achenes was collected meadows affected by the sea winds, between halophy- from La Azohı´a (Cartagena, Murcia; 378330800N; tic thyme bushes. It was first classified as Endangered 181002200W; altitude 30 m). This area has a semi-arid and later as Critically Endangered (Sa´nchez et al., 2004) Mediterranean climate characterized by irregular according to the International Union for Conser- rainfall and a harsh, dry summer period. Annual vation of Nature categories. It is currently protected mean precipitation is around 300 mm, and mean by a regional law of Murcia (BORM, 2003). annual temperature is 178C. August is the warmest Despite the degree of threat to this species, there month, with an average temperature of 24.98C and a are few references to its distribution and the plant maximum of 428C. The coldest month is January, with communities in which it appears (Sa´nchez et al., an average temperature of 10.68C and the minimum 2004), and no reports on its biology and ecology. always .08C. Central fruit heads at similar states of In general, in order to establish appropriate measures maturation were harvested in July 2009 from .400 for the conservation of a species, it is necessary to plants. Achenes were separated according to the white determine its reproductive biology and ecology. One or dark type using a stereoscopic microscope of the most important aspects of the reproductive (Olympus SZ61), and stored for 6–7 d at room biology of a species, especially in arid and unpredict- temperature before germination studies. able environments, is the response of seed germination (Gutterman, 1993). Since the existence of hetero- morphic achenes is not mentioned in the botanical Morphological characterization of achenes and description of A. chrysantha (Tutin et al., 1980), the aims embryos of the present study were to: (1) explore morphological and anatomical variability in achenes; (2) identify Using a stereoscopic microscope with a micrometer, possible differences in dormancy and germination the length, width and pappus length were determined behaviour; and (3) determine possible adaptive for 50 randomly selected achenes of each type. After advantages of this variation in A. chrysantha achenes. measuring of the achenes, the pericarp was removed to determine the length and width of the embryos. Similarly, to estimate the mass of the achenes and Materials and methods embryos, 50 achenes of each type were weighed using a Mettler Toledo XP56 electronic microbalance (with Plant material 0.001 mg precision). The capitula (fruit heads hereafter) of A. chrysantha Achene germination tests (with yellow flowers on a rather convex disc of 12–25 mm in diameter), have peduncles up to 6 cm in Effect of temperature and light on the germination length. The receptacle is hemispherical to oblong– of fresh achenes ovoid, and rounded at the apex (Tutin et al., 1980). Each fruit head contains about 100–130 achenes, with To determine the influence of light and temperature on receptacular bracts between them, arranged in several achene germination, 100-achene lots of each type were rows (7–12) on the disc. Achenes are obconical, incubated at each of the following constant tempera- generally shorter than 2 mm, with ten granular ribs ture regimes: 10, 15, 20 and 258C, and an alternating Morphology and germination of Anthemis chrysantha temperature regime at 128C in darkness and 208Cin 12/208C. Embryo germination was checked every light. Each 100-achene lot was distributed into four 2–3 d for 37 d. To determine if the hardness of the dark samples of 25 achenes. Each sample was incubated in a achene pericarp could affect embryo germination, four 9-cm-diameter Petri dish, on a double layer of filter samples of 25 dark achenes with their fruit coats were paper moistened with 4 ml of distilled water, in scarified slightly at the basal end (radicle end) or at germination chambers (Sanyo MLR-351H, Osaka, the apical end (cotyledon emergence point) and Japan) with a temperature and light control system incubated in light at 12/208C for 37 d.
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