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Development, Fatty Acid Composition, and Storage of Drupes and Seeds from the Endangered Pondberry (Lindera Melissifolia)

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Development, Fatty Acid Composition, and Storage of Drupes and Seeds from the Endangered Pondberry (Lindera Melissifolia) BIOLOGICAL CONSERVATION I37 (2007) 489-496 , . ..... 2. available at www.sciencedirect.com ,---@ *-:" ScienceDirect ELSEVIER journal homepage: www.elsevier.com/locate/biocon Development, fatty acid composition, and storage of drupes and seeds from the endangered pondberry (Lindera melissifolia) Kristina Connofl*, Gretchen ~chaefeP,Jillian Donahooc, Margaret ~euall~,Ernile ~ardinefl, 'ITocy kIawkinsd, Dan wilsond, Nathan ~chiff, Paul ~arnel~,Ted ~einingefl aUS Forest Service, Restoring and Managing Longleaf Pine Ecosystems, 520 Devall Dr., Auburn, AL 36849, United States bl~lWeyerhaeuser Road, Aiken, SC 29801, United States '6302 Lange Drive, Colorado Springs, CO 80918, United States d~~ Forest Service, Center for Bottomland Hardwoods Research, P.O. Box 227, Stoneville, MS 38776, United States ARTICLE INFO ABSTRACT Article history: Pondberry (Lindera melissifolia [Walt.] Blume: Lauraceae) is an endangered, dioecious, clonal Received 9 January 2007 shrub that grows in bottomland hardwood forests in the southeastern United States. hior Accepted 12 March 2007 work has emphasized vegetative reproduction associated with the clonal nature of this spe- Available online 26 April 2007 cies. Little has been published about the early morphological and biochemical characteris- tics of the fruit as they mature. Fruits, drupes originating from the axillaty buds, were Keywords: collected evety 30 days after anthesis and examined for seed structure development and Pondbeny fatty acid composition of the fruit and seed. Sixty days after anthesis, fruits had not formed Seed development an organized embryo/cotyledon, weighed 0.1 i 0.001 g, and measured 7.1 * 0.04 mm x Fatty acids 4.3 * 0.03 mm. Ninety days after anthesis, a complete seed had formed within the drupe. Seed storage Of the total drupe weight (average0.23 i 0.01 g), the seed comprised 33% of the mass gained from 60 days after anthesis. Overall composition of the seed and pulp lipids changed signif- icantly over the course of development. Myristic, palmitic, steric, oleic, linoleic, and linolenic fatty acids were revealed by the lipid analyses. Lauric acid was not found in any of the early seed lipid samples but it increased in quantity as seed matured to become the dominant fatty acid in this tissue. Conversely, pulp contained only small amounts of lauric acid; its fatty acid profile was dominated by oleic acid. Fully hydrated seeds stored well for 16 months at both 4 "C and -2 "C. Although drying had a deleterious effect on germination when dried seeds were conventionally stored at 4 OC, seeds that had been dried for 24 h to a moisture content of 8.6% were successfully stored in liquid nitrogen. Published by Elsevier Ltd. 1. Introduction rare in occurrence (Steyermark, 1949; Radford et al., 1968; Kral, 1983). It grows in bottomland hardwood forests in six Pondberly (Lindera melissifolia [Walt.] Blume: Lauraceae) is a states in the southeastern United States and has recently federally listed endangered shrub that may always have been been rediscovered in a seventh. Though the species is found * Corresponding author: Tel.: +I334 826 8700;fax: +I334 821 0037. E-mail addresses: kconnort3fs.fed.u~ (K. Connor), Gretchen.schaeferQweyerhaeuser.com (G. Schaefer), [email protected] (J. Donahoo), mdevall63fs.fed.us (M. Devall), [email protected] (E. Gardiner), [email protected] (T. Hawkins), dwilson0263fs. fed.us (D. Wilson), nschifft3fs.fed.u~(N. Schiff), phamel63fs.fed.u~(P. Hamel), tleininger62fs.fed.u~(T. Leininger). 0006-3207/$- see front matter Published by Elsevier Ltd. doi:10.1016/j.biocon.2007.03.011 490 BIOLOGICAL CONSERVA? over a relatively broad range, it is distributed sparsely within Cryopresemation protocols have been developed for this temtory, occumng in largely isolated localities. The 1- some seeds that prove difficult to store (Towill and Bajaj, 2 m tall plants are usually found in clonal colonies, a result 2002; Engelmann, 2003). Techniques used include two-step of rhizomatous sprouting. Colonies are often found in areas freezing (Withers and King, 1980), encapsulation-dehydra- that commonly flood in the spring but which are frequently tion (Dereuddre et al., 1991; Shibli and Al-Juboory, 2000), ra- dry in the fall (Devall and Schiff, 2004). pid cooling (Mycock et al., 1995), flash drying (Berjak et al., Reproductive information on the species has concenmated 1989; Wesley-Smith et al., 1992), and vitrification (Touchell on its clonal regenerative capabilities, but it is misleading to et al., 2002). Researchers have been successful in prese~ng consider pondbeny only from this aspect. While vegetative embryonic axes and somatic and zygotic embryos (Shibli reproduction serves to increase colony size, sexual reproduc- and A1-Juboory, 2000; Towill and Bajaj, 2002; Fang et al., tion and seed dispersal are necessary for establishment of 2004). Alternatively, cryopreservation of shoot tips, buds, new colonies. Smith et al. (2004) suggested hydrochory and and apical meristems has also proven successful for many zoochory as potential seed dispersal mechanism in wetland species (Towill et al., 2004). While encouraging, these forests, noting that the showy, persistent fruit of pondbeny labor-intensive processes are primarily undertaken for valu- may favor dispersal by birds or other animals. However, their able agricultural species, fruit trees, or economically impor- research has yet to document an animal dispersal agent capa- tant forest tree species. Additionally, the cryopreservation ble of disseminating seed relatively long-distances to suitable of shoot tips and buds requires destruction of entire plants but uncolonized locales. or plant parts, generally unacceptable when working The sexual reproductive potential of pondbeny is consid- with endangered species except under extreme erable. In Mississippi, flowering typically begins in February, circumstances. before leaves emerge. The dioecious plants produce small Our purpose in this work was to examine (1) the morpho- yellow flowers on axillary inflorescences. Though late frosts logical development of pondbeny fruit and seeds; (2) fatty can damage or destroy flower and seed crops, colonies of fe- acid composition of the storage lipids of seeds and fruit pulp male plants often produce an abundant crop of bright red, as development progresses and (3) maturation and storability spicy-scented drupes. Indeed, in some years individual fe- of the seeds. Such information will provide basic life history male stems can contain in excess of 250 drupes (Connor; per- and biochemical information about the species and can be sonal observation), representing a large energy investment used to evaluate the potential for establishing artificial germ- for the plants. However, little else is known of the species' plasm reserves. population ecology and, thus far, its basic fruiting biology. Fundamental information about pondberry fruit develop- 2. Materials and methods ment, fruit biochemistry, and seed maturation and storability is unknown. ?tnro studies were initiated to address the objectives. The first Plant species conservation often depends on the storabil- investigated drupe development, from pollination through ity of its seeds (Touchell and Dixon, 1993). Stored seeds or maturity. In addition, we tracked the fatty acid biochemistry plant tissues can provide a safety net for species found in of drupes as they developed. The second study determined low numbers in the wild and can be used as a source for rein- the storability of pondbeny seeds and suggested a successful troduction if a population fails (Touchell and Dixon, 1993;Wil- seed handling protocol. kinson et al., 2003). The majority of seeds from temperate forest species are 'orthodox' (Roberts, 1973), meaning they 2.1. Study site can be dried to moisture contents of less than 12%.They be- come metabolically quiescent when dried and are conven- Plant colonies studied for seed development were located on tionally stored at temperatures ranging from 4" to -18 "C or the Delta National Forest, Sharkey County, Mississippi, in cryostored in liquid nitrogen at -196 OC. However, there are the Yazoo River Basin of the Lower Mississippi Alluvial Val- seeds that are desiccation-sensitive. Sub-orthodox (or inter- ley, USA. The site supports a typical bottomland hardwood mediate) and recalcitrant seeds show sensitivity to drying forest cover for the area. Overstory trees are primarily and/or cold temperatures (Roberts, 1973; Ellis et al., 1990). If sweetgum (Liquidambar styraciflua L.), sugarbeny (Celtis lae- carefully handled, sub-orthodox seeds can retain viability vigata Willd.), and red maple (Acer rubrum L.). Nuttall oak and still be dried to as low as 10-12% moisture content. Recal- (Quercus nuttallii E.J. Palmer) and boxelder (Acer negundo L.) citrant seeds, however, must be stored fully hydrated; thus also contribute to the canopy but are of lesser importance both they and fungi found on their outer surfaces are meta- (Hawkins et al., 2004). Climate in the area is typical of the bolically active. Metabolically active seeds can continue to humid, subtropical region of the Northern Temperate Zone. mature and germinate while in storage, and loss of recalci- The growing season averages 229 days long, with 27 "C as trant seeds to fungal infection is not uncommon. These pro- the average temperature in July and 7.5 "C as the average cesses serve to drastically shorten any useful period of seed January temperature (Scott and Carter, 1962; Scott et al., storage and complicate germplasm
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