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Pre-Inoculation by an Arbuscular Mycorrhizal Fungus Enhances Male Reproductive Output of Cucurbita Foetidissima

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Pre-Inoculation by an Arbuscular Mycorrhizal Fungus Enhances Male Reproductive Output of Cucurbita Foetidissima Int. J. Plant Sci. 161(4):683±689. 2000. Copyright is not claimed for this article. PRE-INOCULATION BY AN ARBUSCULAR MYCORRHIZAL FUNGUS ENHANCES MALE REPRODUCTIVE OUTPUT OF CUCURBITA FOETIDISSIMA Rosemary L. Pendleton1 USDA Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, 735 North 500 East, Provo, Utah 84606, U.S.A. Male and female reproductive output of Cucurbita foetidissima, a gynodioecious native perennial, was examined in a 2-yr greenhouse/outplanting study. Plants were divided into three treatment groups: (1) a low- phosphorus (P) soil mix control; (2) a low-P soil mix with the addition of mycorrhizal inoculum (Glomus intraradices); and (3) a high-P soil mix. Plants were outplanted after one summer of greenhouse growth and harvested in the fall of the second year. High-P treatment plants grew best during the ®rst year, having signi®cantly longer vines than either low-P treatment. By the end of the second year, however, treatment had no signi®cant effect on either aboveground biomass or weight of the tuberous storage root. Tissue concen- trations of N and P also did not differ signi®cantly with treatment. Male reproductive output was signi®cantly enhanced by the addition of mycorrhizal inoculum, resulting in a threefold increase over control plants in the production of male ¯owers. In contrast, treatment had no signi®cant effect on aspects of female reproductive output, including number of female ¯owers, percent fruit set, total fruit biomass produced by the plant, or mean fruit weight. Fruit production was correlated with vegetative aboveground biomass and is likely re¯ective of carbon status. These results suggest that mycorrhizal colonization may differentially in¯uence male and female components of plant reproduction. Keywords: buffalo gourd, Cucurbita foetidissima, Glomus intraradices, male reproductive output, gynodioecy, sex allocation. The majority of terrestrial vascular plant species form some may play a signi®cant role in reproductive success of both type of mycorrhizal association, those with arbuscular my- sexual functions. The effect of the soil microbial environment corrhizal fungi (AMF) being the most common (Gerdemann on ¯oral sex expression remains largely unexplored. However, 1975). The association of mycorrhizal fungi with a host plant recent publications report a signi®cant increase in pollen grain root is known to aid in the uptake of phosphorus and certain number, size, and vigor of mycorrhizal zucchini (Stephenson other elements from the soil, often resulting in an increase in et al. 1994, 1998; Lau et al. 1995). plant growth and tissue nutrient content (Mosse 1973; Rhodes The objective of this study was to determine the effect of and Gerdemann 1980; Stribley 1987). Additional bene®ts in- inoculation with an arbuscular mycorrhizal fungus on male clude reduced susceptibility to pathogens and improved resis- and female components of reproduction, including ¯oral sex tance to salinity and drought (Hirrell and Gerdemann 1980; expression, using a native perennial, Cucurbita foetidissima. Rosendahl and Rosendahl 1990, 1991; Ruiz-Lozano et al. This species was chosen for two reasons. As a native, C. foe- 1995; Harrison 1997; Stahl et al. 1998). While studies dem- tidissima has not been subject to intense horticultural selection onstrating increased growth due to mycorrhizal colonization for response to added nutrients and should be re¯ective of abound, the role of mycorrhizae in plant reproduction has other native ®eld species. Also, the use of a monoecious per- received far less attention. Mycorrhizal studies in which male ennial allowed for easy assessment of male and female repro- and female reproductive components were examined sepa- ductive output. rately are rare. Certain plants, including cucurbits, are known to alter their ¯oral sex expression in response to environmental conditions, Material and Methods including light intensity, photoperiod, temperature extremes, soil moisture and nutrient availability, and trauma to leaves, Cucurbita foetidissima HBK, or buffalo gourd, is a long- buds, or storage tissue (Galun 1961; Freeman et al. 1980; lived perennial common to wash bottoms and roadsides in Condon and Gilbert 1988; Delesalle 1989, 1992 and references arid regions of the western United States and northern Mexico therein). Soil phosphorus and nitrogen levels have been shown (Bemis and Whitaker 1969). Populations are gynodioecious, to affect both male and female reproductive success in Cu- composed of both monoecious and gynoecious individuals, curbita pepo (Lau and Stephenson 1993, 1994). These ®ndings presumably controlled by a single locus (Dossey et al. 1981; suggest that colonization by arbuscular mycorrhizal fungi, Kohn 1989). Plants are robust, having a large ¯eshy storage which is known to affect uptake of both of these nutrients, root of up to 40 kg and numerous stems that range from 6 to 12 m in length (Hogan and Bemis 1983; Kohn 1989). Repro- 1 E-mail [email protected]. duction is both sexual, by means of seed, and asexual, through Manuscript received August 1999; revised manuscript received January 2000. adventitious rooting at the nodes. Buffalo gourd has been in- 683 684 INTERNATIONAL JOURNAL OF PLANT SCIENCES vestigated as a potential dryland food and fuel crop (Hogan m between plants. Pots were placed in the ground and the root and Bemis 1983; DeVeaux and Shultz 1985). systems allowed to extend into the surrounding soil. Soil at Seeds of C. foetidissima were collected from plants growing the site had 2.2% organic matter, with a pH of 7.5, 23 ppm near the Snow®eld exit of I-15 in Washington County, Utah. available N, 17 ppm available P, 171 ppm available K, and Seeds were surface-sterilized by shaking in 70% ethanol for 1 an electrical conductivity of 0.93 dS/m (bicarbonate extraction; min and 2% bleach for 15 min, followed by rinsing in sterilized Page et al. 1982). Watering was accomplished by means of a deionized water. Seeds were transferred to sterile blotter papers drip irrigation system, and the area surrounding each plant in plastic petri dishes and strati®ed at 27C for 2 wk before was covered with weed cloth and gravel to prevent adventitious being transferred to a 307C germinator. All seeds germinated rooting at the nodes. within 3 d. Flowering was monitored on a regular basis throughout the Pregerminated seeds were planted in 15-cm pots containing second growing season. The presence of male and female ¯ow- one of three soil treatments: (1) a low-P potting mixture (low- ers was marked at each node using colored tape. At the end P); (2) a low-P potting mixture to which spores of Glomus of the growing season, stem lengths were measured and a map intraradices in an expanded clay carrier (Nutri-link H1000, of ¯owering nodes constructed for each plant. Plants that pro- Native Plants) had been added (low-P1AMF); and (3) a high- duced only female ¯owers were classi®ed as gynoecious. Plants P potting mixture (high-P). A total of 20 pots per soil treatment having at least one male ¯ower were classi®ed as monoecious. were planted with two to three seeds each, then thinned to Root samples were collected from soil layers directly below one plant per pot on emergence of the germinating seeds. The each pot and examined for mycorrhizal colonization. System- base potting mixture for all treatments was composed of peat atic sampling of the entire root system was not possible, how- moss, turface, vermiculite, and sand in a 4:2:3:2 ratio, ever, because of the great depth to which these roots grow. amended with fertilizer at a rate of 700 g dolomite, 350 g Vines, gourds, and storage tubers were harvested, dried, and limestone, 95 g calcium nitrate, 294 g gypsum, 126 g Os- weighed. Vine leaf and stem tissue from 10 randomly chosen mocote 18-6-12 (manufactured by Grace Sierra), 65 g STEM plants per treatment were ground and analyzed for content of trace minerals, and 4 g Fe-138 per 0.0832 m3 of potting mix- N, P, K, Zn, Fe, Mn, Cu, Ca, and Mg by the Plant and Soil ture. Superphosphate was added to the high-phosphorus mix- Testing Laboratory at Brigham Young University. Tissue ni- ture at a rate of 64 g per 83.25 L of potting soil. The mixtures trogen was determined using the Kjeldahl procedure (Horwitz were steamed at 717C for 1 h. Following steaming, water ex- 1980). Content of other bioessential elements was determined tracts (Warncke and Krauskopf 1983) of the mixture had a using atomic absorption procedures on tissue samples digested pH of 6.75, with 123 ppm NO3-N and 126 ppm plant-avail- in a 1 : 5 solution of concentrated sulfuric and nitric acid (Hor- able K. Plant-available P was 0.6 ppm for the low-P mixture witz 1980). and 10.6 ppm for the high-P mixture. Soil analyses were car- Biomass and colonization data were statistically analyzed ried out at the Brigham Young University Plant and Soil Testing using the GLM and CORR procedures on SAS, version 6.1, Laboratory, Provo, Utah. for the personal computer (SAS 1989). Mean separations were Plants were grown in the greenhouse for 8 wk, after which accomplished using the Student-Newman-Keuls multiple- 20 plants of each treatment were transplanted into 26-L con- range test. Differences in plant tissue concentrations were an- tainers ®lled with the appropriate low- or high-P potting mix- alyzed using MANOVA. Percentage data were arcsine trans- ture described above. A spot check of root samples taken at formed before analysis. Differences in the numbers of male the time of transplanting con®rmed that mycorrhizal coloni- and female ¯owers per plant were examined using the non- zation had taken place in plants of the low-P1AMF treatment parametric Kruskal-Wallace test, followed by the Dunn Q-test (Koske and Gemma 1989).
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