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Techniques for Breaking Seed Dormancy of Rainforest Species from Genus Acronychia

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Techniques for Breaking Seed Dormancy of Rainforest Species from Genus Acronychia Liyanage, Offord and Sommerville (2020). Seed Science and Technology, 48, 2, 159-165. https://doi.org/10.15258/sst.2020.48.2.03 Research Note Techniques for breaking seed dormancy of rainforest species from genus Acronychia Ganesha S. Liyanage*, Catherine A. Offord and Karen D. Sommerville The Australian PlantBank, The Royal Botanic Gardens and Domain Trust, Mount Annan, NSW 2567, Australia *Author for correspondence (E-mail: [email protected]) (Submitted February 2020; Accepted March 2020; Published online April 2020) Abstract We tested for dormancy in three species of Acronychia (Rutaceae) occurring in the rainforest in eastern Australia, A. imperforata, A. laevis and A. oblongifolia, by incubating fresh intact seeds on 0.8% water agar for one month at 25/10°C. Four different techniques were then tested for their effect on dormancy: (i) incubation of intact seeds on agar incorporating gibberellic acid (GA3); (ii) seed coat removal (decoating); (iii) scarification near the radicle emergence point (scarification-emergence point); and (iv) scarification opposite the radicle emergence point (scarification-back). Imbibition tests were performed to determine whether dormancy was due to an impermeable seed coat. Germination differed among treatments, but all three species showed a similar pattern. Intact seeds showed < 6% germination after one month indicating the presence of dormancy. Highest germination (> 65%) was observed following scarification-emergence point treatment. Seed coat removal also resulted in increased germination (40-47%), in comparison with intact seeds, but GA3 and scarification-back treatments did not (< 12%). Though the seedcoats of all species were permeable, increased germination responses to decoating and scarification-emergence point treatments suggest scarification is required to clear the radicle emergence point. This may be a useful dormancy-breaking technique for Acronychia spp. and may be suitable for related Rutaceae species. Keywords: Acronychia, dormancy-breaking, Rutaceae, scarification, seed biology, seedcoat Experimental and discussion Seed dormancy is a trait that prevents seed germination during unfavourable environmental conditions where subsequent seedling establishment is likely to be unsuccessful (Baskin and Baskin, 2014). Ecologically, this is a beneficial trait that spreads the risk of failure of seedling establishment over time in disturbance-prone environments (Philippi and © 2020 Liyanage et al. This is an open access article distributed in accordance with the Creative Commons Attribution Non Commercial (CC BY-NC 4.0) license, which permits others to distribute, remix, adapt, build upon this work non-commercially, and license their derivative works on different terms, provided the original work is properly cited and the use is non-commercial. See: https://creativecommons.org/licenses/by-nc/4.0 159 GANESHA S. LIYANAGE, CATHERINE A. OFFORD AND KAREN D. SOMMERVILLE Seger, 1989; Philippi, 1993). However, from a conservation perspective, seed dormancy constrains the implementation of ex situ seed banking and in situ plant restoration. Good seed germination is essential to the success of both conservation practices. For example, standard seed banking involves drying seeds to 3-7% moisture content and storing them in subzero (≤ −20°C) freezer conditions (CBD, 2012). Testing how species respond to this drying and freezing involves a series of germination tests. In addition, germination tests need to be performed regularly to assess the viability of seed bank collections over time. Performing those tests is a challenge when the seeds are difficult or slow to germinate. Plant species from the Rutaceae family have often been recorded as producing seeds that are difficult to germinate due to dormancy (Auld, 2001; Floyd, 2008), yet studies on dormancy breaking techniques for Rutaceae species of non-sclerophyllous ecosystems, such as rainforests, are very limited (Elliot and Jones, 1982; Martyn et al., 2009). The genus Acronychia J.R. Forst. and G. Forst. is one of 152 genera in the Rutaceae family (POWO, 2019) and nearly 20 Acronychia species are found in Australian rainforests (Atlas of Living Australia, 2020). This is a genus that contains nationally endangered species such as A. littoralis T.G. Hartley and J.B. Williams (Environment Protection and Biodiversity Conservation Act, 1999) and that occurs in habitats that are themselves under threat (Sommerville et al., 2018). The development of ex situ seed bank collections and techniques to relieve dormancy is therefore urgent. In this current study we investigated three Acronychia species occurring in Australian rainforest, A. imperforata F. Muell., A. oblongifolia (A. Cunn. ex Hook.) Endl. Ex Heynh and A. laevis J.R. Forst. and G. Forst., in order to understand the treatments required to break dormancy and facilitate germination. This knowledge will contribute to efforts to conserve the species in ex situ seed bank collections. Acronychia imperforata is an endemic species found in lowland and littoral rainforests. Acronychia oblongifolia and A. laevis are recorded as occurring in warmer rainforests and dry and subtropical rainforests respectively (Floyd, 2008). For each species, mature fruits were collected from more than three well-established plants growing in The Australian Botanic Garden, Mount Annan, Australia (34°03'22.3''S 150°46'29.0''E). Acronychia oblongifolia fruits were collected in May 2018; A. imperforata and A. laevis fruits were collected in August 2018. Seeds were extracted from the fruits by hand and healthy- looking seeds were selected for experiments that commenced within two weeks of collection. As low seed fill is commonly reported in Rutaceae species, seed fill was first assessed using 40-60 seeds per species (depending on seed availability). Seeds were dissected and observed under a microscope and scored as ‘filled’ if they contained a firm green/white endosperm and embryo. The presence of dormancy was tested by placing three replicates of 15 seeds per species on Petri dishes containing 0.8% water agar. The Petri dishes were sealed with cling wrap to control moisture loss and the seeds were incubated at alternating temperatures of 25/10°C with a 12/12 hour light/dark cycle. The incubation temperature represented the maximum and minimum temperatures recorded around the time of seed dispersal for the regions in which these species mainly occur. Germination was recorded every second day for one month; a seed was considered to have germinated when the emerging radicle was > 1 mm in length. At the end of the incubation period, ungerminated 160 DORMANCY BREAKING OF ACRONYCHIA seeds were checked for firmness by pressing the seed with forceps. Any remaining firm seeds were dissected to check for the presence of a green/white embryo and endosperm and assess whether the seeds remained viable. The final germination percentage was adjusted for empty/unfilled seeds and an adjusted germination < 50% was considered to indicate dormancy (Baskin and Baskin, 2014). Four dormancy-breaking treatments were then applied to each species: (i) incubation of intact seeds on agar incorporating gibberellic acid (GA3 treatment); (ii) full seed coat removal (de-coating); (iii) scarification near the radicle emergence point (scarification- emergence point); (iv) scarification opposite the radicle emergence point (scarification- back). Three replicates of 15 seeds per treatment per species were placed on either 0.8% water agar (treatments ii, iii and iv) or 0.8% water agar incorporating 0.25% gibberellic acid (treatment i). An additional three replicates of 15 intact seeds were placed on 0.8% water agar as a control. Treatment (i) was not performed for A. laevis due to low seed numbers. All Petri dishes were incubated and seeds monitored for germination as described above. An imbibition test was performed to determine whether seedcoat permeability may have a role in maintaining dormancy. Two samples of 10 manually scarified and intact seeds were weighed then placed on 0.8% water agar and re-weighed at 24 hour intervals until all the scarified seeds had undergone imbibition. The germination percentages of intact seeds were calculated for each species to determine the presence/absence of dormancy. Differences in germination among dormancy- breaking treatments for each species were compared using a one factor Generalised Linear Model with a binomial error structure and logit link function. Where germination responses were significantly different, Tukey’s tests were performed to compare differences among treatments (Hothorn et al., 2008). Differences in seed mass between intact and scarified seeds resulting from imbibition were compared using a t-test. All data were analysed using the R 3.5.1 statistical platform (R Core Team, 2018). Seed fill was > 87% for both A. imperforata and A. oblongifolia and 60% for A. laevis. Final viable seed numbers were, therefore, corrected for empty seeds before the comparison of germination data for A. laevis. Intact seeds incubated on water agar germinated poorly for all species (5.0, 5.6 and 1.7% for A. imperforata, A. laevis and A. oblongifolia, respectively; figure 1). Application of dormancy-breaking treatments produced a similar germination pattern for all three species, with significant differences among treatments (A. imperforata df = 4, χ2 = 82.7, P < 0.001; A. laevis df = 2, Z = 29.4, P < 0.001; A. oblongifolia df = 4, χ2 = 119.3, P < 0.001). Gibberellic acid had no effect on germination
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