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Ectomycorrhizal Formation by Micropropagated Clones of Eucalyptus Marginata Inoculated Vs^Ith Isolates of Pisolithus Tinctorius

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Ectomycorrhizal Formation by Micropropagated Clones of Eucalyptus Marginata Inoculated Vs^Ith Isolates of Pisolithus Tinctorius AVir I'hytnL (19X9). 111. 20')-214 Ectomycorrhizal formation by micropropagated clones of Eucalyptus marginata inoculated vs^ith isolates of Pisolithus tinctorius BY C. M. TONKIN', N. :\/[ ALAJ CZU K- AND J. A. McCOMB' ' School of Environmental and Life Sciences, Murdoch University, Mttrdoch, Western Australia 6150 *-Division of Forest Research CSIRO, Private Bag, P.O. Wembley, Western Australia 6014 {Received 15 April 1988; accepted 19 October 1988) S U M M A R ^' /tiicalvpliis iiHuiiiiiala Donn ex Sm. and I'isolilhiis liiicloriiis (Pcrs.) Col; aiul CoLich were co-cultLircd to obtain ec-tomycorrhiziil formation /)( vitro. One isolate of P. tincloriiis formed mycorrhizas with aseptic seedlings of a ju^•enile clone deri\-ed from a 4-month-old seedling, and lour clones derived from crowns of mature trees. A second P. liiirhiriiis isolate formed mycorrhizas with only the clones from mature trees. Successful combinations resulted in formation of a mantle followeil b\- a MartiK net and epidermal eell elongation. The ftmgal/seedlings or fungal/seedling clone combinations which did not produce ectomycorrhizal roots, were characterized by a mantle but lacked a Hartig net, and formed an abundance of polyphenols throughout the root, (jenotype, maturity and fungal specificity are key faetors influencing successful ectom\ corrhizal formation on E. marginata by P. tiiictoiiiis in vilro. Key words: Kctotn>'corrhiza, micropropagation, Pisolilliiis liiirlornis. Kiualyplus iiuux'iiiala, clone. 1982; Piche, Peterson & Acherley, 1983; Massicotte, I N'lHODl'C TI ON Peterson & Ashford, 1987a, 6). Ectomycorrhizal IVlicropropagation of forest trees showing attributes formation is rapid with typical mantling of the root for superior firowth ma\- pro\icle the means ol surface and formation of a Hartig net. To date, increasing prodtictix'itx' in reafforestation pro- however, most studies on ectomycorrhizal synthesis gramtnes. Significatit increases in growth from sel- in vitro have utilized genetically variable seedlings ected clones of pine and hardwoods have in part or a single clonal line derived from a seedling been attribtited to improved eHicienc\' in nutrient (Malajczuk et al., 1982; Malajczuk & Hartney, uptake and retranslocation (Libby & Rauter, 19<S4; 1986). Sheppard & Cannell, 1975). This paper reports on the interaction between the Mycorrhizas are characteristic of all species of plant genotype and the fungal isolate in formation of forest trees (Trappe, 1962, 1977) and aid in tiutrient ectomycorrhizas in vitro. The tree species used was (particularly phosphortis) uptake (Malajczuk, jarrah (Eucalyptus marginata Donn e.x Sm.) which McComb & Lonei-agan, 1975). A range of soil and was chosen as it is an importatit forest species in plant factors are known to affect tnxcorrhizal root Western Australia and there is a wide range of clonal development, iticluding plant genot> pe and strain ol material available (Bennett & McComb, 1982). The fungal symbiont (Mason, 1975). To maximize the ability of five clones to form mycorrhizas with two benefits of micropropagation it will be necessary to isolates of P. tinctorius was examined. ensure that these plants form effecti\'e mycorrhizas. Pisolilliiis tinctorius (Pers.) Cok and Cotich is a M.^TERI.ALS .AND MliTHODS species which forms ectomycorrhizas /;; vitro with Production of axeiiic jarrah seedlings several tree genera (Alalajczuk, Molina .& Trappe, * Address for ofliirints. Seeds of jarrah (code no. D393, Department of 210 M. Tonkin, N. Malnjcziik and J. A. MrCoinb Table 1. Clonal Itttes of Eucalyptus marfi;inata tised for examiiting ectontycorrhizal formation in vitro Clonal line No. Source of explant Plant details Cr of mature tree Survivor on a site with dieback (Phytophthora ciniianwmi) Karnet, W. Australia 6 Crown of mature ti"ee Leafmincr (Perthida fih'phopa) resistant tree, Shentoii Park, W. Australia 7 Crown of mature tree Leafminer {Perthida glyphopa) resistant tree, Shenton Park, W. Australia 20 Shoots from 4-month-old Survivor of a P. cinnamomi and seedling waterlogging pot trial, Murdoch University, W. Australia 53 Crown ol mature tree Survivor on a site with dieback (P. cinnamomi) Karnet, W. Australia Conservation and Land Management, Western growtb (> 5 mtn d ') on ERM medium (witbout Australia) were surface sterilized in 95"',, ethanol hormones). for 30 s followed by 2"o sodium bypochlorite for 20 min and then rinsed tbree times in sterile water. Seed coats were removed and embryos germinated In vitro uioculatioit aseptically on moist filter paper and incubated in darkness at 25 °C. After 10 d, germinated seeds were Ten-day-old rooted plantlets of tbe five clonal lines transferred to 8 x 2-5 cm polycarbonate tubes con- of jarrab, and 1 0 d-old jarrah seedlings were placed taining 10 ml of half strength JVIurasbige & Skoog into polycarbonate tubes containing 10 ml ERM (1962) medium without hormones solidified with witbout hormones, and inoculated with an agar plug 0-8 "„ agar. Time zero was the day of germination. Irom a vigorously growing ectomycorrbizal fungus. Tubes were then incubated at 25 °C under a 16 b light (150//mol m"'-s"')/8 h dark cycle. Plantlets Production of axenic clonal plant lets of jarrah and seedlings were examined 3 and 6 weeks alter Shoot cultures of jarrah clonal lines were obtained inoculation for formation of ectomycorrhizas. In from Mr 1. Hennett, Murdoch University, Western addition after 25 d, plantlets of clonal line 7 and Australia (H^ible I) and were maintained on eucalypt seedlings were sampled and the number of leaves, multiplication medium (Bennett & McComb, 1982). root tips and ectomycorrbizal roots, and root lengtb p\)ur clones were derived from tbe crowns of mature were recorded. trees and are referred to as 'mature clones'. One clone from a 4-month-old seedling is referred to as a 'juvenile clone'. Cultures were divided and trans- Ectontycorrhizat anatomy ferred to fresh media every three weeks and Samples of the main (tap) and lateral roots were incubated at 25 °C under ligbt (50/imolm "s '). excised from clonal lines 7, 20, 53 and seedlings at 5 Individual shoots of selected clonal lines were and 18 days after inoculation. Root pieces were fixed transferred to eucalypt root initiation medium in 3 "/(, glutaraldebyde in 0-025 M phosphate buffer (F.RM) and incubated at 25 °C for root induction (pH 7-3) on ice for 2 h, then overnight under vacuum (Hennett & McComb, 1982). Cultures were initially infiltration at room temperature. Tissues were rinsed placed in darkness for 8 d, then a 16 b light (15()/( twice in buffer and post-fixed in 2",, osmium mol m - s ')/8 h dark regime. Time zero was tbe day tetroxide in ()-O25 M phosphate buffer for 1-2 h at of root initiation. room temperature. Samples were then washed in distilled water and dehydrated through an acetone Fungal i.tolates series, 10, 25, 50, 75 and 90 "o for 1 5 min followed by two 15 min rinses of 100",, acetone. Tissue was Two isolates of Pisolithus tinctorius were used. H98 infiltrated in an acetone: Spurr's series 3:1,2:1, 1:1, was isolated from basidiocarps associated with jarrah 1:2, 1:3 (eacb for 48b), rinsed in 100",, Spurr's and marri (/t. calophylla R. Iir.) growing on coastal (Spurr, 1969) and finally embedded in 100 "v, Spurr's sandy soil tiear Eagle Hay National Park, WA in at 70 °C for 24 b. Sections 1-2 //,M thick were cut on 1981. H323 was isolated from basidiocarps associated a Porter Hlum JH-4 microtome and examined after with jarrab and marri growing on lateritic soil at staining with methylene blue (1 "/„ in 1 "/„ sodium Nannup, WA, in 1983. Hoth isolates show rapid tetraborate) or they were cleared in saturated as of iitiii'()f>r<>f>/ii>at('d l<"ucal\'pti.is marginata 211 Table 2. In vitro de^'etopnient of ectontycorrhisas on with the fungus (i.e. main roots as well as tine roots) jarralt phnttlcts aitd seedliitj^s .J iveeks after iitoctttatiott formed cctomycorrliizas. The.se roots were generalK' zvitlt isolates of Fisolithus tinctorius ('.J replicates per simple and unbranched, lacked root hairs and liad treatment) larger diameters in comparison to non-mycorrhizal roots. The mantle was yellow-brow n witli numerous Jarrah P. tinctorius hyphae radiating outwards (Fig. 1). clonal Control line H98 H323 (no fungus) Effect of P. tinctorius iiioctthitiou oit plattt Ljiotctlt 3 6 Both P. tiitctoritts isolates retarded shoot grow tli of 7 micropropagated plantlets as assessed b\' number ot 2O lea\es. In seedlings onh' isolate H')8 reduced shoot 53 growth (Table 3). Seedlings Although it was difficult to determine a time zero that would be comparable for plantlets and seedlings, + , Pre.sence of ectomycorrhizal roots on each replicate. —, Absence of ectomycorrhizal roots on each replicate. both liad roots initiated at time zero and were then One mycorrhizal tip observed on one plant. grown on basal media for 10 d before inoculation. Seedlings had a slower rate of root dexelopment, with fewer laterals, compared to plantlets of the same potassium hydroxide (in ethanol) for 1-2 min, rinsed age. Fewer ectomycorrhizal root tips tbrmed on in water, and then examined unstained or stained seedlings compared to plantlets. Root growth (botli with methylene blue. length and number of tips), was sc\ erely reduced in inoculated plantlets and seedlings when compared to the uninoculated controls. The formation of eeto- R I-: .s IIL T s myeorrhizas resulted in larger mean lateral root Formation of eetomyeorrhizas diameters of plantlets and seedlings inoculated witli T'hree weeks after inoculation with P. tinetoritis both /-". liiteloritts isolates. isolate H98, eetomyeorrhizas were observed on seedlings, the juvenile clone and all the mature Effect of P. tinctorius ittoctitatioit oit root anatomy clones. P. tiitrloritis isolate H323 formed eetomyeor- rhizas only on clonal lines derived from mature In uninoculated elonal lines and seedlings pol\- trees (Table 2).
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