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After-Ripening, Stratification, and Perigynia Removal Enhance

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After-Ripening, Stratification, and Perigynia Removal Enhance Physiological dormancy is com- After-ripening, Stratification, and Perigynia mon in the Cyperaceae and may be Removal Enhance Pennsylvania overcome or reduced by one or more of the following treatments: 1) after- Sedge Germination ripening (dry storage of seeds under ambient temperatures before sowing), 1 2) GA3, and 3) cold stratification Esther E. McGinnis and Mary H. Meyer (Baskin and Baskin, 1998, 2004). Broom sedge (Carex scoparia) germi- ADDITIONAL INDEX WORDS. Carex pensylvanica, dormancy, propagation, nation was enhanced by up to 2 years Cyperaceae, achene of after-ripening (Larson and Stearns, 1990). Elongated sedge (Carex elongata) SUMMARY.Pennsylvaniasedge(Carex pensylvanica) has horticultural and restoration and remote sedge (Carex remota)in- potential, but the achenes are difficult to germinate due to complex dormancy creased germination following after- requirements. This study identified treatments to overcome physiological dormancy and determined light and temperature requirements for optimum germination. ripening in comparison with fresh We first tested the effects of perigynia removal and light on achene germination. In the achenes (Schutz, 1997b). However, second experiment, achenes were subjected to varying durations of dry-cold or some wetland Carex species had dry-warm storage conditions and a presowing soak in gibberellic acid (GA3). In higher germination percentages when a third experiment, we studied whether storage conditions, cold stratification, and stored cold and moist (Budelsky and sowing temperatures affected germination. Pennsylvania sedge germination was Galatowitsch, 1999). improved by dry-warm storage, perigynia removal, cold stratification, and Although GA3 failed to stimulate germination in light. germination in black and white sedge (Carex albonigra), ebony sedge [Carex ebenea (Amen and Bonde, ennsylvania sedge is commonly published, and native plant nurseries 1964)], and hood’s sedge [Carex used for forest restoration (Mottl propagate plants by division. Over- hoodii (McDonough, 1969)], it has Pet al., 2006) and has horticul- coming dormancy and understanding been shown to increase germination tural potential as a shade-tolerant germination requirements are essen- for other monocots such as sand rye- groundcover and low-maintenance tial for economically propagating grass [Leymus arenarius (Greipsson, lawn species (Darke, 2007). This her- pennsylvania sedge on a commercial 2001)], green needlegrass [Stipa vir- baceous perennial is native to dry de- basis. idula (Fulbright et al., 1983)], east- ciduous forests of the eastern half of Few Carex species exhibit physi- ern gamagrass [Tripsacum dactyloides temperate North America (Gleason cal dormancy or other germination (Rogis et al., 2004)], and four Aus- and Cronquist, 1991). It produces barriers as a result of their unique tralian grass species (Hagon, 1976). attractive slender leaves that form morphology. Carex are distinguished In contrast to GA3 pretreatment, cold a 12-inch mound of foliage that ex- from other genera within the Cyper- stratification has been shown to success- pands through long and short rhi- aceae by a bladder-like sac called the fully alleviate physiological dormancy zomes (Bernard, 1990) to form mats perigynium (perigynia, plural) that in many Carex species (Hoag et al., of 3 m2 (Mottl et al., 2006). Unlike tightly adheres to the hard pericarp 2001; Kettenring and Galatowitsch, most lawn species, it thrives in dry of the achene (Amen and Bonde, 2007a, b; Schutz and Rave, 1999). partial shade and is uniquely suited to 1964). The perigynium prevents ger- Although the most effective stratifi- the competitive environment under mination in nebraska sedge (Carex cation temperatures for Carex species large trees. Pennsylvania sedge also nebrascensis) and northwest territory can vary, temperatures below 12 °C provides spring interest because it sedge (Carex utriculata) (Hoag et al., are most effective (Brandel and Schutz, blooms in mid-April to mid-May in 2001; Jones et al., 2004). In other 2005). Optimum stratification dura- southern Ontario and in the northern cases, Carex species respond to tradi- tion may range from 0.5 to 6 months United States (Crins and Ball, 1983). tional physical dormancy treatments for Carex species (Kettenring and Achenes ripen and dehisce in June in such as acidic scarification (Ishikawa Galatowitsch, 2007a). Minnesota (Table 1). Difficulties in et al., 1993) or pericarp nicking Pennsylvania sedge may have ad- achene germination limit the use of (Amen and Bonde, 1964). It is un- ditional germination requirements. A pennsylvania sedge for large horti- known whether pennsylvania sedge light requirement enables woodland cultural and restoration projects. exhibits physical dormancy or other Carex species to take advantage of No germination protocol has been germination barriers. gaps in leaf litter on the forest floor Department of Horticultural Science, University of Minnesota, 305 Alderman Hall, 1970 Folwell Ave- nue, St. Paul, MN 55108 Units We thank Alan Wade from Prairie Moon Nursery, To convert U.S. to SI, To convert SI to U.S., Winona County, MN, for donating achenes and plant multiply by U.S. unit SI unit multiply by material for this experiment. 0.0929 ft2 m2 10.7639 Mention of a trademark, proprietary product, or 3.7854 gal L 0.2642 vendor does not imply endorsement by the University 2.54 inch(es) cm 0.3937 of Minnesota or its approval to the exclusion of other 16.3871 inch3 cm3 0.0610 suitable products or vendors. 1 ppm mgÁL–1 1 1Corresponding author. E-mail: [email protected]. (°F – 32) O 1.8 °F °C(1.8·C) + 32 • April 2011 21(2) 187 RESEARCH REPORTS Table 1. Origin and collection month of ripe pennsylvania sedge achenes used in University of Minnesota germination experiments, 2005 and 2006. Location Address Collection mo. Prairie Moon Nursery Section 15, Township 105N, Range 7W, June 2005, 2006 Winona County, MN University of Minnesota, Section 17, Township 116N, Range 23W, June 2006 Landscape Arboretum Carver County, MN Lebanon Hills Regional Park Sections 34 and 35, Township 27N, June 2006 Range 19W, Dakota County, MN Cedar Creek Natural History Area Section 34, Township 34N, Range 23W, June 2006 Anoka County, MN or in the deciduous forest canopy required for the experiments; there- achenes were placed on top of the (Vellend et al., 2000). Germination fore, mature achenes were collected sand in each dish. The sand was of Carex species increased when im- from three additional locations in moistened with deionized water, and bibed achenes were exposed to white Minnesota (Table 1). In both years, more water was added as needed. fluorescent light (Kettenring et al., filled achenes were separated from Four petri dishes (four replications) 2006; Schutz and Rave, 1999). In a unfilled achenes using an air column for a total of 60 achenes were assigned groundcover study, the frequency of separator and tested for viability using to each treatment. All four petri pennsylvania sedge decreased under a 1% solution of 2,3,5-triphenyltetra- dishes were kept in a clear plastic tree canopy compared with clearings zolium chloride as described in Grabe bag (1 gal) to prevent excessive evap- (Collins and Good, 1987). Optimum (1970). Viability percentages ranged oration. The entire experiment was germination temperature has not been from 77% to 81% in both years. repeated 1 week later. identified for pennsylvania sedge. Eu- Because of the similarity of viability, Achenes exposed to white light ropean temperate sedges typically re- 2006 achenes were combined. The were placed in a growth chamber quire warm temperatures (>28 °C) for total number of achenes per petri dish (Environmental Growth Chambers, germination (Grimes et al., 1981). was multiplied by the estimated via- Chagrin Falls, OH) and subjected to Schutz and Rave (1999) reported that bility percentage, and this number 200 mmolÁm–2Ás–1 of irradiance under a diurnally fluctuating temperature was deemed to be the maximum cool white fluorescent bulbs (model regime also enhances germination of possible germination per dish. F72T12/CW/VHO; Philips Light- some Carex species because this mimics EXPT.1:PERIGYNIA REMOVAL ing, Somerset, NJ) for 12 h per day, spring temperatures. Alternating day AND LIGHT EXPOSURE. This experi- with alternating 26 °C, 10-h days/ and night temperatures of 27 and ment was conducted in 2006 with 18 °C, 10-h nights with a 2-h transition 15 °C increased germination for un- achenes that had been stored dry at period between temperatures. Achenes stratified achenes for the majority of 22 °C, 50% relative humidity (RH) with no light exposure were placed in 12 temperate North American wet- for 8 weeks. The achenes were divided the same growth chamber, but petri land Carex species (Kettenring and for the following four treatments: 1) dishes were double wrapped in alumi- Galatowitsch, 2007a). intact perigynia, no light; 2) intact num foil to prevent light penetration. The objectives of this project perigynia, exposure to white light; 3) Germination counts were conducted were to test whether pennsylvania removed perigynia, no light; and 4) daily or every second day for 8 weeks. sedge germination is enhanced by removed perigynia, exposure to white Germinated seedlings were periodi- treatments designed to overcome light. Perigynia were removed by cally removed. Counts for the dark germination barriers posed by the rubbing the achenes in a hand trough treatments were made in a dark room perigynia and by the physiological between two layers of hard dimpled
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