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Sclerocarya Birrea (A. Rich.) Hochst. Subsp. Caffra (Sond.) Kokwaro

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Sclerocarya Birrea (A. Rich.) Hochst. Subsp. Caffra (Sond.) Kokwaro SEED TECHNOLOGY HORTSCIENCE 44(1):119–124. 2009. and Goyvaerts, 2004). As industrial demand for its products is increasing, there is a growing concern about the sustainable supply After-ripening, Light Conditions, (Nwonwu, 2006) and conservation of wild populations. Thus, in the quest for the domes- and Cold Stratification Influence tication and improvement of S. birrea, under- standing of seed germination of this plant is Germination of Marula [Sclerocarya essential. Several members of Anacardia- ceae, including Sclerocarya, are character- ized by a drupe fruit with a stony endocarp. birrea (A. Rich.) Hochst. subsp. The endocarp of Sclerocarya and other related genera has a specialized structure, caffra (Sond.) Kokwaro] Seeds the operculum, through which the germinat- Mack Moyo, Manoj G. Kulkarni, Jeffrey F. Finnie, and ing embryo emerges (Von Teichman and 1 Robbertse, 1986). Hills (1933) stated that Johannes Van Staden the Anacardiaceae exhibits some remarkable Research Centre for Plant Growth and Development, School of Biological seed protection mechanisms by means of a and Conservation Sciences, University of KwaZulu-Natal, Pietermaritzburg, hard lignified endocarp and, intriguingly, the Private Bag X01, Scottsville 3209, South Africa most ingenious devices to allow emergence of the germinating embryos. This ingenious Additional index words. seed storage, photoinhibition, light intensity opening device (operculum) represents one of the most sophisticated opening mecha- Abstract. Marula [Sclerocarya birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro nisms in the germination of seeds (Von (Anacardiaceae)] is used in many African countries as a food crop and is also in demand Teichman and Robbertse, 1986). Typical of for industrial purposes. The fruit pulp has high vitamin C levels and the nuts have a high the Anacardiaceae, the germinating unit protein and oil content. The fruit pulp is commercially used in the production of an (seed) in S. birrea is the true seed plus alcoholic beverage (Amarula Cream) and the oil is gaining importance in the cosmetic endocarp (Game´ne´ et al., 2004; Li et al., industry. Although attempts are being made to domesticate this high-value indigenous 1999). tree, there is very limited information available on aspects of seed germination. Our Although considerable research has been study investigated the role of light, temperature, cold stratification, and after-ripening on undertaken on this species, there is still a seed germination of S. birrea. Temperatures between 25 and 35 8C favored germination dearth of knowledge on some aspects of its of opercula-removed seeds under continuous dark conditions. White light completely seed biology. Game´ne´ et al. (2004) inconclu- inhibited seed germination with the inhibitory effect being reversed when seeds were sively suggested that seed after-ripening, a transferred to dark conditions. This photoinhibitory effect on opercula-removed seeds decrease in mechanical resistance of the was lost after 12 months of seed storage at room temperature in the dark. Cold operculum after storage, or a combination stratification (5 8C) of intact seeds for 14 days significantly improved germination (65%) of both factors can improve germination of S. as compared with nonstratified seeds (32%). Pregermination treatments (acid scarifica- birrea seeds. Another research gap relates to tion, boiling water, dry heat, soaking, and plant growth regulators) of S. birrea seeds did the effect of light on the germination process not promote germination. Seeds of S. birrea can be considered orthodox because they (Von Teichman et al., 1986). Generally, tolerated desiccation without significant loss of viability. Both intact and opercula- under natural conditions, temperature, light, removed seeds readily imbibe water suggesting physiological rather than physical water, oxygen, and mechanical pressures are dormancy. The highest germination percentage was recorded under constant dark some of the important factors that can influ- conditions at 25 8C for opercula-removed seeds exposed to an after-ripening period of 12 ence seed germination of species like S. months. This study indicates that after-ripening, light conditions, and cold stratification birrea. The purpose of this study was to are critical factors for germination of S. birrea seeds. identify the possible environmental and mechanical cues influencing the germination of S. birrea seeds. Sclerocarya birrea is native to the semi- chemical analyses indicate that the kernels arid deciduous savannas of sub-Saharan have higher protein and oil content than most Materials and Methods Africa (Muok et al., 2007) and is one of the of the popular nuts, including walnuts, hazel- most highly valued indigenous trees in south- nuts, chestnuts, and almonds (Wynberg et al., Seed collection. Fruits of Sclerocarya ern Africa (Von Teichman and Robbertse, 2003). birrea were collected in Feb. 2007 from the 1986). Besides a number of medicinal uses Humankind has benefitted from S. birrea Mpumalanga Province of South Africa. (Eloff, 2001), the importance of S. birrea is as a source of nutrition for more than 10,000 Fruits were depulped and cleaned as underpinned by its diverse characteristics years (Nwonwu, 2006). Apart from contrib- described in the seed leaflet of the Danida such as high levels of vitamin C and protein, uting to rural diets, the fruit is used to brew an Forest Seed Centre (2003). The seeds (endo- quality stable oil, and the novel flavor of its alcoholic beverage with an annual gross carps) were separated from the pulp, washed, fruit. The vitamin C content of S. birrea fruit value of $80 to $100 U.S. per household dried, and stored in brown paper bags at room juice is approximately four to five times more (Emanuel et al., 2005; Shackleton et al., temperature (22 ± 2 °C) for 4 weeks before than the levels found in the average 2008). The oil is in high demand in the phar- being tested for germination ability. Seeds orange juice (Jaenicke and Thiong’o, 2000; maceutical and cosmetic industries (Kleiman used to determine the initial moisture content Mojeremane and Tshwenyane, 2004). Bio- et al., 2008; Nwonwu, 2006), whereas the were not stored. nuts are used in the food industry for making Seed germination. Before each germina- a range of products, including chocolates. tion test, seeds were surface-decontaminated With the realization of its market value, there by soaking for 15 min in 0.5% (w/v) solution Received for publication 18 Sept. 2008. Accepted is notable growth in the trade of S. birrea of mercuric chloride (HgCl ). Subsequently, for publication 25 Sept. 2008. 2 The National Research Foundation (NRF), Pretoria products stimulated by local and industrial seeds were thoroughly rinsed under tap and and the University of KwaZulu-Natal are thanked demand (Emanuel et al., 2005). then distilled water. Before the germination for financial support. Owing to its economic potential, S. birrea experiments, seeds were soaked for 24 h in 1To whom reprint requests should be addressed; has been earmarked for crop development the dark at room temperature (22 ± 2 °C) in e-mail [email protected] and improvement in southern Africa (Mollel distilled water (covering 75% of the seed) for HORTSCIENCE VOL. 44(1) FEBRUARY 2009 119 the hard endocarps to imbibe water. Seed Moisture content determination. True 4-h intervals for 96 h. Percentage water germination was carried out on cotton wool seeds (embryonic axis and cotyledons) (Fig. uptake was calculated on the basis of actual moistened with distilled water and placed 1) were excised from the stony endocarp of increase in seed mass over the initial seed in plastic containers (10.5 · 10.5 · 13.5 cm) marula seeds and moisture content was deter- mass (Hidayati et al., 2001): in growth chambers equipped with cool mined gravimetrically by weighing before white fluorescent lamps (Osram L 58W/640, and after oven-drying at 110 °C for 48 to 96 h %W s = ½ðW i – W dÞ=W d 3 100 Mu¨nchen, Germany) emitting a photosyn- until a constant weight was obtained. Mois- thetic photon flux density (PPFD)of100 ture content was calculated on the basis of where Ws = increase in mass of seeds; Wi = mmolÁm–2Ás–1 over a wavelength band of 400 fresh weight: (% moisture content) = (fresh mass of seeds after a given interval of to 700 nm. Light intensity was measured with weight – dry weight)/fresh weight · 100. The imbibition; and Wd = initial mass of seeds. a quantum radiation sensor (Model Skp 215; results represent the means of the moisture Effect of temperature. Intact seeds were Skye Instruments Ltd., Llandrindod Wells, content values of 20 embryos (± SE) obtained soaked in distilled water for 24 h before Powys, UK). Seeds were considered to have from two separate experiments. exposing them to different temperatures. germinated when the radicle had emerged at Water uptake. Water uptake was deter- Seed germination was determined for both least 2 mm (Bewley, 1997). Each treatment mined using both intact (Fig. 1) and opercula- intact and opercula-removed seeds under consisted of 25 seeds and was replicated four removed (Fig. 1) seeds that had been stored at alternating light (16-h photoperiod of 100 times. All the experiments were repeated ambient room temperature in the dark for 12 mmolÁm–2Ás–1) and continuous dark conditions twice. Germination was recorded daily. The months. Intact and opercula-removed seeds (containers were wrapped with aluminum seeds that were subjected to continuous dark (25 seeds per replicate) were placed on cotton foil). The seeds were incubated at constant conditions were examined under a ‘‘green safe wool moistened with distilled water in plastic temperatures of 10, 15, 20, 25, 30, 35, and 40 light’’ (wavelength of 510 nm and PPFD of containers (10.5 · 10.5 · 13.5 cm) and °C and alternating temperature of 30/15 °C 0.2 mmolÁm–2Ás–1) in the dark (Kulkarni et al., incubated under cool fluorescent white light (14 h/10 h) in plant growth chambers (Con- 2006).
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