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Water Absorption and Dormancy-Breaking Requirements of Physically Dormant Seeds of Schizolobium Parahyba (Fabaceae – Caesalpinioideae)

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Water Absorption and Dormancy-Breaking Requirements of Physically Dormant Seeds of Schizolobium Parahyba (Fabaceae – Caesalpinioideae) Seed Science Research, page 1 of 8 doi:10.1017/S0960258512000013 q Cambridge University Press 2012 Water absorption and dormancy-breaking requirements of physically dormant seeds of Schizolobium parahyba (Fabaceae – Caesalpinioideae) Thaysi Ventura de Souza, Caroline Heinig Voltolini, Marisa Santos and Maria Terezinha Silveira Paulilo* Departamento de Botaˆnica, Universidade Federal de Santa Catarina, Floriano´polis 88040-900, Brazil (Received 14 July 2011; accepted after revision 18 January 2012) Abstract seeds have been a source of controversy (Fenner and Thompson, 2005; Finch-Savage and Leubner-Metzger, Physical dormancy refers to seeds that are water 2006). A definition of dormancy that has been impermeable. Within the Fabaceae, the structure proposed recently is that dormancy is an innate seed associated with the breaking of dormancy is usually property determined by genetics that defines the the lens. This study verified the role of the lens in environmental conditions in which the seed is able to physical dormancy of seeds of Schizolobium para- germinate (Finch-Savage and Leubner-Metzger, 2006). hyba, a gap species of Fabaceae from the Atlantic Five classes of seed dormancy are recognized, and one Forest of Brazil. The lens in S. parahyba seeds of them is physical dormancy (Baskin and Baskin, appeared as a subtle depression near the hilum and 2004), which is caused by a seed (or fruit) coat that opposite the micropyle. After treatment of the seeds prevents absorption of water (Morrison et al., 1998; with hot water, the lens detached from the coat. Baskin and Baskin, 2001; Smith et al., 2002). Blocking water from contacting the lens inhibited water Physical dormancy is known to occur in 17 families absorption in hot-water-treated seeds. High constant (308C) and alternating (20/308C) temperatures pro- of angiosperms, including the Fabaceae (Baskin and moted the breaking of physical dormancy and Baskin, 2000; Funes and Venier, 2006), where it occurs in germination in non-scarified seeds. Maximum percen- many species. Water-impermeability of the coat (or in tage of germination occurred earlier for seeds some species the fruit coat) is caused by the presence of incubated at 20/308C than for those incubated at one or more layers of elongated, lignified Malpighian 308C. Seeds with a blocked lens did not germinate at cells that are tightly packed together and impregnated alternating or high temperatures. This study suggests with water-repellant chemicals (Morrison et al., 1998; that alternating temperatures are probably the cause Baskin and Baskin, 2001; Smith et al., 2002; Baskin, of physical dormancy break of seeds of S. parahyba in 2003). Under natural conditions, it has been suggested gaps in the forest. that physical dormancy is not broken by seeds passing through the digestive tracts of an animal or by cracks in the coat caused by animals (Baskin and Baskin, 2001; Keywords: Fabaceae, lens, physical dormancy, water Fenner and Thompson, 2005). One characteristic that uptake suggests this hypothesis is correct is the presence of a specialized anatomical region in physically dormant seeds that develops an opening where water can enter Introduction the seeds (Baskin and Baskin, 2001). Several types of specialized structures (‘water gaps’) have been found in For plants, it is important that seed germination occurs 12 of the 17 families that have physical dormancy; for in the right place and at the right time, and, for this example, the carpellary micropyle in Anacardiaceae; reason, most species have mechanisms that delay the bixoide chalazal plug in Bixaceae, Cistaceae, germination, such as seed dormancy (Fenner and Cochlospermaceae, Dipterocarpaceae and Sarcolaena- Thompson, 2005). The definitions of dormancy in ceae; the imbibition lid in Cannaceae; the chalazal plug in Malvaceae; the lens and hilar slit in Fabaceae (Baskin et al., 2000) and the micropyle-water gap complex in *Correspondence Geraniaceae (Gama-Arachchige et al., 2011). However, Email: [email protected] in some Fabaceae (subfamilies Caesalpinioideae and 2 T.V. de Souza et al. Mimosoideae) the lens is absent (Gunn, 1984, 1991) and 1981; Matheus and Lopes, 2007), the seed coats were after treating some legume seeds to break physical made impermeable in four ways: (1) extrahilar region dormancy, cracks develop in the extrahilar region blocked with paraffin; (2) hilar region blocked with or in the hilum that permit entrance of water into the paraffin; (3) hilum blocked with Super Bonderw glue seeds (Hu et al., 2008, 2009). (Henkel, Jundiai, Brazil); and (4) lens blocked with Several artificial techniques are used to break physical Super Bonderw glue. A control group was of non- dormancy in seeds, including mechanical, thermal and dormant, non-blocked seeds. Twenty seeds were chemical scarification, enzymes, dry storage, percussion, utilized for each treatment. Seeds were placed in low temperatures, radiation and high atmospheric transparent plastic boxes of 11 £ 11 £ 3.5 cm on two pressures (Baskin and Baskin, 2001). Studies on seeds layers of filter paper (Whatman No. 1, Whatman with physical dormancy have contributed greatly to International Ltd, Maidstone, England) with 10 ml of our understanding of water gaps, the effects of various distilled water. The boxes were stored at 208C with a factors (e.g. drying, heating, low temperatures and photoperiod of 12 h/12 h. Incubated seeds were alternating temperatures) in breaking physical dor- counted at intervals of 2 or 3 d for 19 d, during which mancy under natural conditions, and the rate and path time germination was observed. of water entrance into seeds that have become permeable (Baskin and Baskin, 2001). Under natural conditions, it is Analysis of seed coat features known that temperature is an important environmental factor for breaking physical dormancy in seeds (Baskin The hilar regions of five intact and five thermally and Baskin, 2001). Va´zquez-Yanez and Orozco-Segovia scarified seeds were fixed in 2.5% glutaraldehyde (1982) verified that the highly fluctuating temperature in a 0.1 M sodium phosphate buffer at pH 7.2 and that occurs in gaps, but not in forest understorey, breaks dehydrated in a graded ethanol series. Sections of physical dormancy in gap forest species. 40 mm thickness were cut using a sliding microtome. Schizolobium parahyba (Fabaceae–Caesalpinioideae) Histochemical tests were made utilizing Sudan IV for is a pioneer woody species from the Atlantic Forest of suberin, cutin, oils and waxes; acid phloroglucinol Brazil that occurs mostly in gaps and along forest and iron chloride for lignin (Costa, 1982); and toluidine borders, with physically dormant seeds and anemo- blue for polychromatic reactions to lignin and cellulose choric seed dispersal (Carvalho, 2003). The imperme- (O’Brien et al., 1965). Images were taken with a digital able seed coat of this species can be broken artificially camera connected to an optical microscope (Leica MPS by boiling water or mechanical scarification (Caˆndido 30 DMLS). For scanning electron microscopy (SEM) et al., 1981; Freire et al., 2007; Matheus and Lopes, 2007). analyses, the dehydrated pieces of five intact and five The aim of this work was to study the seeds of scarified seeds were immersed in hexamethyldesila- S. parahyba with the objectives of: (1) locating the water sane (HMDS) for 30 min, as a substitute for critical gap in the seeds; (2) describing the anatomical point drying (Bozzola and Russell, 1991) and then structure of the water gap; and (3) testing the effect mounted on aluminium stubs and blocked with a of alternating temperatures on breaking the physical gold layer (40 nm thick). The pieces were viewed using dormancy of the seeds. a Jeol JSM 6390 LV scanning electron microscope. To verify the presence of callose in the seeds, sections of non-fixed samples of the hilar and extrahilar Materials and methods regions of five intact seeds were immersed in 0.05% aniline blue with a 0.1 M potassium phosphate buffer Seed collection at pH 8.3 (Ruzin, 1951). As a control, some sections were immersed only in the potassium phosphate buffer. The Seeds of S. parahyba, which remain enclosed in the sections were observed using an Olympus BX41 similar-shaped papery envelope of endocarp resembling microscope, with a mercury vapour lamp (HBO 100) a wing, were collected from the ground soon after wind and a blue epifluorescence filter (UMWU2), at 330– dispersal, during spring, in a section of Atlantic Forest 385 nm excitation and 420 nm emission wavelengths. located in the municipality of Florianopolis, Santa Images were taken with a Q-imaging digital camera 0 00 0 00 Catarina, Brazil (27835 36 S, 48835 60 W). The endocarp (3.3 mpixel QColor3C) and the software Q-captures was removed, and the seeds were stored in plastic bottles Pro 5.1 (Q Images, Surrey, British Columbia, Canada). at room temperature until they were used. Effect of alternating temperatures on germination Location of the water entrance region and dormancy break After artificially breaking dormancy of the seeds by Seeds were immersed in 5% sodium hypochlorite for placing them in water at 988C for 1 min (Caˆndido et al., 5 min and then washed three times in distilled water. Physical dormancy in seeds of Schizolobium 3 For some of the seeds, the region with the lens was with the blocked lens was only 1.0%. However, 50% of covered using Super Bondw glue, which made the the scarified seeds with only the hilum blocked seeds impermeable. Then the seeds were placed in germinated (Fig. 1). The germination levels at the last transparent plastic boxes on a 5 cm autoclaved layer of day of incubation were similar for seeds blocked in the sand moistened with distilled water. The boxes were lens and in the hilar regions, but significantly different stored at 208C, 308C and a 12 h/12 h alternating for scarified seeds and scarified seeds blocked in the temperature regime of 20/308C with a photoperiod of extrahilar region and hilum (P # 0.05).
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