Beneficial Role of Arbuscular Mycorrhizal Fungi on Florida Native
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PALMS Fisher & Jayachandran: Mycorrhiza Vol. 52(3) 2008 JACK B. FISHER Beneficial Fairchild Tropical Botanic Garden, 11935 Old Cutler Road, Role of Miami, Florida 33156 USA (address for correspondence: [email protected]) Arbuscular and Dept. of Biological Sciences, Florida International Mycorrhizal University, Miami, Florida 33199 USA Fungi on AND K. JAYACHANDRAN Florida Department of Environmental Studies and Southeast Environmental Research Native Center, Florida International University, Palms Miami, Florida 33199 USA Mycorrhizas, the symbiotic associations of roots and soil fungi, are present in Florida native palms collected in the wild. However, the role of mycorrhizas in the growth of these palms is unclear. We studied seedlings of four palms native to the Everglades to see how mycorrhizas affect growth and uptake of phosphorus. Our findings will be helpful in proposed Everglades restoration projects. The symbiotic fungi are called arbuscular microscope. Within the root cells, the fungus mycorrhizal fungi (AMF) which are common forms tiny tree-like structures, called and found in the majority of land plants arbuscules, which are the places where including cycads, conifers (except the pine nutrients and water are exchanged between family), and most trees and crop plants, the two organisms. The fungus cannot make including palms. The fungi penetrate and grow its own food and receives organic nutrients inside the roots but are not visible on the root (sugars and amino acids) from the surface with the naked eye. They also do not photosynthetic plant. In return, the plant form large, obvious reproductive bodies (i.e., receives mineral nutrients and water through mushrooms or truffles). Therefore, fungal the fungi. Smith and Read (1997) and Wang presence can only be verified after clearing, and Qiu (2006) gave extensive reviews about staining and then observing the roots with a the general biology of mycorrhizas. PALMS 52(3): 113–123 113 PALMS Fisher & Jayachandran: Mycorrhiza Vol. 52(3) 2008 1. Representative plants of Coccothrinax, plants unpotted and washed free of soil. Treatments: 1 = no AMF, no microbes; 2 = no AMF, added microbes; 3 = live AMF, added microbes; 4 = no microbes, added 10 mgkg-1 P. Although no comprehensive surveys of the (African oil, coconut and date) and many wild palm family have been conducted, AMF have palm species (reviewed in Fisher & been found in every species of palm that has Jayachandran 1999). Since this last review, been studied. The first well documented AMF were reported in wild Desmoncus palms description of a mycorrhiza in a palm and its (Ramos-Zapata et al. 2006), wild Florida native beneficial effect on palm growth was given for palms (Fisher & Jayachandran 2005) and old the peach palm (Bactris gasipaes Kunth) by cultivated date palms (Bouamri et al. 2006). Janos (1977). Later, St. John (1988) presented But the mere presence of AMF does not tell us an historical review of AMF in palms. AMF about the benefits of AMF for the plant. occur in all the three main commercial palms Although controlled experiments found that 114 PALMS Fisher & Jayachandran: Mycorrhiza Vol. 52(3) 2008 addition of AMF to potted palms increased Our experimental study has practical growth (reviewed later in detail), there have implications for palms being grown for been no experiments with native Florida restoration and for palms growing naturally palms. Here, we report on several experiments in the Greater Everglades ecosystem. Native with native palms that investigate some of the palms are key species in both wetlands and effects of AMF on palm seedlings that were uplands of pine forest and hardwood grown in pots with native soil. This native hammocks of this ecosystem. We need to sandy soil had low levels of phosphorus and understand the propagation and cultural other nutrients. We want to know if AMF requirements of the native palms better to promote growth and increase the uptake of improve their successful outplanting into phosphorus in these palms, similar to the natural environments. AMF may play an effects of AMF on rare non-palms that were important role in the Everglades, a naturally tested on the same native infertile soil (Fisher low phosphorus (P) environment. Additions & Jayachandran 2002). of P, as from nursery operations, can be 2. AMF in the cortex of ultimate feeder roots of palms. a. Intercellular hyphae and arbuscules in Sabal. b. Intracellular hyphae and arbuscules in Sabal. c. Vesicle and arbuscules in Sabal. d. Intercellular hyphae in Acoelorrhaphe. A, arbuscule; H, hypha; V, vesicle. Scale line = 50 µm. 115 PALMS Fisher & Jayachandran: Mycorrhiza Vol. 52(3) 2008 ecologically disruptive (Fish & Wildlife Service least 12 wk old before use. The inoculum 1999). AMF may be a potential substitute for samples showed heavily colonized root fertilizers in nursery production of plants used fragments and many AMF spores. Spores in restorations if native palms respond to AMF. collected on the 250 µm sieve were mostly the genus Gigaspora or Scutelospora; spores smaller Materials and Methods than 250 µm were mostly Glomus spp. Soil and Plants root fragments were mixed well and used as live inoculum (Treat. 3, below). A part of the In the Greater Everglades ecosystem in same inoculum was steam pasteurized for 2 h subtropical Florida, four species of palms are one time and used as an inoculum control for a visible and ecologically important all treatments without live AMF (Treat. 1, 2 & component of the native plant communities. 4, below). Thus, all treatments received similar Wetlands have the Everglades or Paurotes palm additions of organic and inorganic P found in [Acoelorrhaphe wrightii (Griseb. & H. Wendl.) the inoculum. H. Wendl. ex Becc.] and pine rocklands have the silver palm [Coccothrinax argentata (Jacq.) Experiment L.H. Bailey], the cabbage palm [Sabal palmetto Seeds were collected from wild plants, surfaced (Walter) Lodd. ex Schult. & Schult.f.], and the sterilized with 1.0% sodium hypochlorite saw palmetto [Senenoa repens (W. Bartram) solution for 15 min, rinsed in water, scattered Small]. The last two palms also occur in plant on the surface of inorganic Perlite® medium communities transitional between wetland in plastic pots, and placed under periodic mist and pine rockland. Urban development, watering. Seedlings with one or two green farming and nursery operations have simple leaves (eophylls) were removed from eliminated much of these wetlands and upland the medium, roots trimmed to 4 cm length communities. (to more easily fit into pots), and transplanted Soil into 5 × 18 cm plastic pots (D40 Deepots™, Stuewe & Sons, Inc., http://www.stuewe.com) Soil samples for physicochemical analysis were each filled with 600 g native sandy soil (see collected from a pine rockland site containing description above) that was twice steam wild plants of Coccothrinax, Sabal and Serenoa. pasteurized. Pots, each with one seedling, were The native pine rockland site had shallow grouped by species. Treatments were randomly sandy soils over a bed of oölitic limestone. Soil arranged in frames and grown on benches in analysis (n = 3 at each depth) was carried out a glasshouse under ca. 50% shade. During the by a commercial testing service (A&L Southern growing period, pots were periodically Agricultural Laboratories, Pompano Beach, FL rearranged within and between species groups. 33064). At the 0–5 cm. depth, color was white Cross contamination was prevented by to gray; pH = 6.8–7.3; weak Bray test = 8 mg separation of pots and care in watering. kg-1P. At the 5–20 cm depth, color was white to yellow-orange; pH = 7.3–7.4; weak Bray test Four treatments were: = 4–8 mg kg-1P. Treatment 1: no AMF; no soil microbes; Soil used to fill pots for greenhouse no P experiments was dug from 5–20 cm depths Treatment 2: no AMF; added soil microbes; and mixed. Soil was sieved through 6 mm no P mesh to remove stones and large root fragments. The average soil-solution P was Treatment 3: live AMF; added soil 0.002 mg L-1 (SE = 0.0011, n = 4), as microbes; no P determined with a water extraction method Treatment 4: no AMF; no soil microbes; by Olsen & Summers (1982). The average soil added 10 mg kg-1 P P (weak Bray test) from 0–20 cm depth = 6 mg Each pot received 20 g of either live or dead kg-1. inoculum (sterilized). Soil filtrate (containing soil microbes but not AMF) consisted of 30 ml AMF inoculum per pot of a soil solution derived from 300 g Nurse cultures of mixed native AMF from pine fresh soil shaken in 2 L distilled water, and rockland were maintained on pigeon pea and filtered through Whatman No. 1 filter paper. sudan grass in 4 L pots containing fresh, Phosphorus addition consisted of five unpasteurized soil from the native habitat of treatments on alternating weeks beginning 3 the species of interest. Nurse cultures were at or 4 wk after transplanting. This brought the 116 PALMS Fisher & Jayachandran:FisherMycorrhiza & Jayachandran (revision) Vol. 52(3) Fig 3 2008- 1 Fig. 3 (4 graphs) Acoelorrhaphe Coccothrinax 3.5 3 d b b d 3 d ab 2.5 2.5 2 o o i i t c t a 2 a R R r cd a a cd r o o 1.5 ) ) g g e c ( d ( 1.5 s s s s a a 1 a a M 1 M 0.5 0.5 0 0 1234 1234 Treatment Treatment Sabal Serenoa 3.5 3 b a 3 a 2.5 a a 2.5 c c 2 o o i i t a t a a R 2 a R r a c r o o 1.5 ) ) g g ( ( 1.5 s s s s a d a 1 b M M b b 1 c 0.5 0.5 0 0 1234 1234 Treatment Treatment 3.