EFFECTS OF ENDOPHYTE INFECTION ON THE GROWTH OF LOL IUM PERENN E L. UNDER D ROUGHT STRESS
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1 2002, 26 (5) Ξ Acta Phytoecologica Sinica 3 33 (, ), ( Lolium perenne L. ) ( Neotyphodium lolii),, ; ;, EFFECTS OF ENDOPHYTE INFECTION ON THE GROWTH OF LOL IUM PERENN E L. UNDER D ROUGHT STRESS LIANG Yu 3 CHEN Shi- Ping GAO Yu-Bao 33 and REN An- Zhi ( College of Life Science, Nankai University, Tianjin , China) Abstract Endophytes are fungi that form inapparent infection within leaves and stems of healthy plants. En2 dophytes or endophytic fungi exist widely in the higher plants and are generally considered as mutual symbiotes to the host plants. The present study examined the association of an endophyte ( Neotyphodium lolii) with its host plant, Lolium perenne cv. Pinnacle. Both endophyte status and drought stress level were considered. En2 dophyte-free ( EF) plants were obtained by a mild heat treatment while endophyte- infected ( EI) plants were germinated directly. After transplantation and four weeks of growth in pots, EF and EI plants were subjected to drought stress of different water status in the soil ; soil moisture was maintained at 20 % to 40 %, 40 % to 60 %, 60 % to 80 %, and over 80 % of the field capacity. Leaf extension rate and tiller numbers were mea2 sured during drought stress, and biomass was determined after recovery stage. Our objective was to determine the influence of endophyte infection on growth characteristics and drought resistance of the host plants. Leaf extension of Lolium perenne shifted during drought stress, while no significant superiority was found in EI plants as compared with EF plants. The EI plants tended to produce more tillers than the EF plants dur2 ing drought stress. After severe stress and recovery stage, the EI populations had a significantly higher root/ shoot ratio than the EF populations, while there was no significant difference in total biomass between EI and EF populations. Our results indicated that endophyte infection might enhance the drought resistance of the host plant by enhancing its tiller production during drought stress and its photosynthate allocation to roots between stress periods. Key words Endophyte, Lolium perenne L., Drought stress, Leaf extension rate, Tiller number, Biomass, - -, 80 ( Carroll, 1988 ; Kalburtji & Mosjidis, 1993 ; Siegel,, 1993), (Clay, 1987 ; Hoveland, 1993) (Clay Ξ : : : ( G ) ( ) 3 :, Present address : Research Center for Biodiversity Conservation, Institute of Botany, the Chinese Academy of Sciences, Beijing ibcas. ac. cn 33 Author for correspondence edu. cn
2 & Holah, 1999), (Lolium perenne L. ) ( Endophyte Infected,, EI) ( Endophyte Free, EF),, 1. 3,, ( Lolium perenne L. ), 13 ( 2 cm), , ( Neoty2 ( ) phodium lolii) Pinnacle,, 85 % 1. 1 EI EF, (43 15 min, min, ), ;,, 1, : EI 60 %, EF 2 0 % EI EF cm, 21 cm, 3 cm 7. 2 kg (, :, > 80 % FC ( Field Capacity ) ;,80 % 60 % FC ;, 60 % 40 % FC ;,40 % 20 % FC,, FC, (15 cm) Fig. 1 Changes of environmental factors during drought stress 2 EI, 3 cm ) ( p = ), % 1 kg ( 1) 24,EI EF 12, ; EF, ( p ,0. 099), ( p = ) 46 d, EI EF :, EF EI (, EI 0 % 10 % 60 % 100 % ), EF, ( p, 4, ,0. 165,0. 067) ;,EF EI ( p = ) , EI EF
3 5 : 623 EI Control 1 (cm) Table 1 Cumulative leaf length of the marked plants during drought stress Mild stress Moderate stress Severe stress Mean ab b a ab ab EF ab a ab bc ab :, ( = 0. 05) Data are presented in the format of mean standard error ; same letter denotes non - significant difference while different letters denote a significant difference ( = 0. 05) ( 2 3 Table 2 and Table 3 are the same as Table 1) 2 Fig. 2 Change in leaf blade length of marked plants during drought stress ( 3, 4 The legends of Fig. 3 and Fig. 4 are the same as those of Fig. 2) 3 Logistic Fig. 3 Tiller number of the marked plants of perennial ryegrass during drought stress period and the fit curves of Logistic equation EF( 2) , ( 4) EI EF , EF, EI EF,,,, EI ( p = ) ; EF,,,EI EF, EI EF, EI ( p , ), ( p = ),, EF ; EF, ( 3) ( p ,0. 021),,EI, ; EF,EI EF, EI EF,,EI ( p < 0. 05) ( 2) EI EF ; EF, Logistic ( p = ,0. 001) : N = K/ (1 + e 2rt ), EI K
4 Table 2 Tiller number by the end of drought stress and parameters of Logistic fit curves Tiller number of marked plant by the end of drought stress EI EF Mean Mean Parameters of Logistic equation r K a ab a a a b bc b c c (g) Table 3 Biomass of population after a 4-month recover period (g) EI EF (DW) (DW) Under-ground biomass Above-ground biomass Roots/ shoots ratio ab b a a a a ab ab a b b a c c a ac b ab a bc a abc b b,1992 ;,1992), Latch (1985),EI EF Be2 lesky (1989), Accessions (, ),, 4 Accessions,EF Fig. 4 Increase of total tiller number during drought stress Belesky (1987), (Clone), ( p = ),, EI EF EI EF,,, EI EF EI EF ( p = ),, EI 3,EF 3. 1,EI, ( ), ( Hsiao et al., 1976 ;,1987 ;
5 5 : 625, EI ( ), ( 3), EI EF Chep lick (2000) 13,, Clay (1987), 10 14, EI EF, ; 6, EI EF Latch (1985) EI EF ; Belesky (1987) EI EF, EI, 5, 4 EI EF ; EI EF EF,1 EI EF, West ( 1993), , EI EF, 42 % 55 %,EF, EI EF, EI, EI EF : EI,, EI, EF EF,,, EF, EI EF,, 3. 3 (Siegel, 1993 ; Arachevaleta et al., 1989 ;, Clay, 1990 ; Bacon, 1993), Clay (1987),Bacon Siegel (1988) De Battista (1990a), 2001),, 12 (, 2001a), (,, 2001b) ( Elmi et al., 2000) 18 % 25 %, Porter (1985) ( Balansia epichloe), (IAA) De Battista (1990b), EF, (Amalric et al., 1999 ;, ( Neotyphodium coenophialum) IAA Latch (1985) Amalric, C., H. Sallanon, F. Monnet & A. Hitmi Gas ( GA) exchange and chlorophyll fluorescence in symbiotic and non- sym2, West (1993) biotic ryegrass under water stress. Photosynthetica, 37 : , , Arachevaleta, M., C. W. Bacon, C. S. Hoveland &D. E. Rad2 EI EF ciffe Effect of the tall fescue endophyte on plant response to environmental stress. Agronomy Journal, 81 : , Bacon, C. W. & M. R. Siegel Endophyte parasitism of Cunningham(1988) tall fescue. Journal of Production Agriculture, 1 :
6 Bacon, C. W Abiotic stress tolerances ( moisture, nutri2 ents) and photosynthesis in endophyte-infected tall fescue. Agri2 culture, Ecosystems and Environment, 44 : Belesky, D. P., O. J. Devine, J. E. Pallas Jr. & W. C. Stinger Photosynthetic activity of tall fescue as influenced by a fungal endophyte. Photosynthetica, 21 : Belesky, D. P., W. C. Stinger &N. S. Hill Influence of endophytes and water regime upon tall fescue accessions.. Growth characteristics. Annals of Botany, 63 : Carroll, G Fungal endophytes in stems and leaves : from la2 tent pathogen to mutualistic symbiont. Ecology, 69 : 2 9. Chen, S. P. ( ),Y. B. Gao ( ),Y. Liang( ) & A. Z. Ren( ). 2001a. Effect of endophyte infection on the concentrations of free proline and ABA of leaves of Lolium Endophyte effects on reproduction of a root- knot nematode ( Meloidogyne marylandi) and osmotic adjustment in tall fescue. Grass and Forage Science, 55 : Hoveland, C. S Importance and economic significance of the Acremonium endophytes to performance of animals and grass plant. Agriculture, Ecosystems and Environment, 44 : Hsiao, T. C., E. Acevedo, E. Fereres & D. W. Henderson Water stress, growth, and osmotic adjustment. Philosophi2 cal Transactions of the Royal Society of London Series B-Biologi2 water deficits. Plant Physiology Communications ( cal Sciences, 273 : Jiang, M. Y. ( ) &J. H. Jing ( ) Primary mechanisms of the inhibition of plant cell elongation growth under ), 28 : (in Chinese) perenne L. under water stress. Acta Ecologica Sinica ( Jing, J. H. ( ) & T. C. Hsiao ( ) Effects ), 21 : (in Chinese) Chen, S. P. ( ),Y. B. Gao ( ),Y. Liang( ) & A. Z. Ren ( ). 2001b. Effect of endophyte infection on protective enzyme activities in leaves of Lolium perenne L. under water stress. Chinese Journal of Applied and Environmental Biol2 ogy ( ), 7 : (in Chinese) of water stress and rewatering after water stress on leaf elongation rate of maize. Acta Phytophysiologica Sinica ( ), 13 : (in Chinese) Kalburtji, K. L. & J. A. Mosjidis Effects of Sericea les2 pedeza residues on cool- season grasses. Journal of Range Manage2 ment, 46 : Cheplick, G. P., A. Perera & K. Koulouris Effect of Latch, G. C. M., W. F. Hunt &D. R. Musgrave Endo2 drought on the growth of Lolium perenne genotypes with and with2 out fungal endophytes. Functional Ecology, 14 : Clay, K. &J. Holah Fungal endophyte symbiosis and plant diversity in successional fields. Science, 285 : Clay, K Effects of fungal endophytes on the seed and phytic fungi affect growth of hybrid ryegrasses. New Zealand Jour2 nal of Agricultural Research, 28 : Li, D. Q. ( ), Q. Zou ( ) & B. S. Cheng ( ) Osmotic adjustment and osmotica of wheat cultivars with different drought resistance under soil drought. Acta Phyto2 seedling biology of Lolium perenne and Festuca arundinacea. Oe2 cologia, 73 : physiologica Sinica ( ), 18 : nese) (in Chi2 Clay, K Fungal endophyte of grasses. Annual Review of E2 cology Systematics, 21 : Cunningham, P. J Productivity and persistence of perennial ryegrass at low rainfall sites in southwest Victoria. In : Research Review 1987/ Pastoral Research Institute, Hamilton, Victo2 ria. 41. Liang, Y. ( ), Y. B. Gao ( ), S. P. Chen( ) & A. Z. Ren( ) Effects of endophyte infection on photosynthesis, transpiration and water use efficiency of Lolium perenne L. under drought stress. Acta Phytoecologica Sinica ( ), 25 : (in Chinese) Porter, J. K., C. W. Bacon, H. G. Cutler, R. F. Arrendale & De Battista, J. P., C. W. Bacon, R. Severson, R. D. Plattner J. D. Robbins In vitro auxin production by Balansia &J. H. Bouton. 1990a. Indole acetic acid production by the fungal endophyte of tall fescue. Agronomy Journal, 82 : epichloe. Phytochemistry, 24 : Siegel, M. R Acremonium endophytes : our current state of knowledge and future direction for search. Agriculture, Ecosys2 De Battista, J. P., J. H. Bouton, C. W. Bacon & M. R. tems and Environment, 44 : Siegel. 1990b. Rhizome and herbage production of endophyte- re2 moved tall fescue clones and populations. Agronomy Journal, 82 : West, C. P., E. Izekor, K. E. Turner & A. A. Elmi Endophyte effects on growth and persistence of tall fescue along a water- supply gradient. Agronomy Journal, 85 : Elmi, A. A., C. P. West, R. T. Robbins & T. L. Kirkpatrick. : :
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