2001, 25 (5 537 543 A cta Phytoecolog ica S in ica Ξ Ξ ΞΞ (, 300071 (L olium p erenne L., (N eotyp hod ium lolii ( A crem onium lolii : 1, (E I (EF ;, E I EF ;, E I EF, ; 2, ; 3,, E I EF EFFECTS OF ENDOPHY TE INFECT ION ON PHOTOSY NTHESIS, TRANSP IRAT ION AND W ATER USE EFF IC IENCY OF LOL IUM P ER ENN E L. UND ER D ROUGHT STRESS L IAN G Yu 3 GAO Yu2Bao 3 3 CH EN Sh i2p ing and R EN A n2zh i (Colleg e of L if e S ciences,n ankai U niversity, T ianj in 300071 Abstract N et assim ilation rate, transp iration rate and w ater use efficiency of endophyte2infected (E I and endo2 phyte2free (EF of L olium p erenne w ere determ ined under drough t so as to investigate the ef2 fect of endophyte infection on the ho st p lant. T he physio logical characteristics of L. p erenne depended m uch on drough t levels, and the effects of endophyte infection w ere sign ifican t under p ro longed severe. 1 A t the late stage of severe drough t, the E I assim ilated CO 2 at a significantly h igher rate than the EF. A fter rew atering, how ever, there w ere no significant differences betw een the two 2 T ransp iration rate of the w as mo re affected by drough t than endophyte infec2 tion. 3 O nly at the later stage of severe drough t, w as the w ater use efficiency of the E I h igher than that of the EF. Key words Endophyte, L olium p erenne, N et assim ilation rate, T ransp iration rate, W ater use efficiency, ( (C lay, 1987; D e Battista et 80, (N eotyp hod ium lolii a l., 1990; Hoveland, 1993; R ead & Cam p, 1986; ( A crem on ium lolii R ice et a l., 1990 : Ξ : 2000209204 : 2001205213 : (39870142 (G2000018601 Ξ3 :, 100093 P resent address: Research Center fo r B iodiversity Conservation, Institute of Bo tany, the Ch inese A cadem y of Sciences, Beijing 100093 E2m ail: coo lrain@ns. ibcas. ac. cn ΞΞ 3 3 A utho r fo r co rrespondence E2m ail: ybgao@nankai. edu. cn
538 25 E I EF ( ( 8 kg ( ; = 1 1 20. 37%,, 6. 31%, 1 kg, 24, E I EF (Siegel, 1993 12, 24, 1. 2, 1999 10 5 11 8 (D e 35 d Battista et a l., 1990; C lay, 1987; R ice et a l., : 1990,, EF E I ( (Be2 0 10% 70% 100%, ; lesky et a l., 1987; R ichardson et a l., 1990, 4,,, 2 4 8 3, : > 75% TAW C (To tal A vailab le W ater Capacity,, 75% 50% TAW C, TAW C, 25% 0 TAW C (,,,,,,,,, (L olium p erenne L., 11 5,, 1. 3 50% 25%, C I2301PS (C ID,, CO 2,, 13 30 14 30, 3 d 1 10 7 11 2 11 8 1, 7 00 17 00, 2 h 1. 1 E I EF 1 0. 4 m 0. 4 m 0. 3 m,, 15 W,, (43 15 m in, 57 25 m in (PA R CO 2,,, EF ; ( 1,, E I 1 1. 4,, 1, 2 3 1,, (Λm o l 5 : E I 75%, EF m - 2 (mm o l m - 2, 2% (mm o l m o l - 1, : 1 M icro soft Excel97,,
5 : 539 = {[P (x 1 + P (x 2 ] tg2} P (x 1 P (x 2, t ( s, 7200 (2 h ; = {[E (x 1 + E (x 2 ] tg2} E (x 1 E (x 2 ( 2 A E I EF, t (s, 7200 (2 h ; 2 2. 1 E I 7 00, E I EF ( 1, E I EF = g,, E I 1 F ig. 1 Changes in environm ental facto rs during drought a: 10 7 O ctober 7 b: 11 2 Novem ber 2 c: 11 8 Novem ber 8 d: The w ho le period of drought g A ir temperature L eaf temperature A ir hum idity (RH Pho to synthetic active radiation (PA R CO 2 A ir CO 2 concentration 2 (10 7 F ig. 2 D iurnal changes of net assim ilation rate and transp iration rate of the trial in the early stage of drought E I E I contro l E I E Im ild EF EF m ild E I E I severe EF EF contro l EF EF severe 3 4 5 T he legends of F ig. 3, F ig. 4 and F ig. 5 are the sam e as tho se in F ig. 2
540 25, E I EF EF 1 (10 7 Table 1 values of diurnal pho to synthesis, diurnal transp iration and w ater use efficiency of L olium p erenne in the early stage of drought Physio logical characteristics of E I and EF T rial value of W ater use value of diurnal diurnal efficiency pho to synthesis transp iration (mmo l CO 2 (mmo l CO 2 (mo m - 2 l H 2O m - 2 mo l - 1 H 2O 253. 3 67. 89 a 167. 52 3. 80 a 1. 541 0. 061 a E I M ild 308. 06 43. 26a 175. 08 7. 76 a 1. 740 0. 189 a 331. 50 73. 85 a 168. 46 5. 41 a 1. 997 0. 503 a 290. 40 78. 47 a 183. 66 11. 43 a 1. 544 0. 345 a EF M ild 236. 52 26. 41a 164. 92 5. 10 a 1. 456 0. 190 a 262. 06 56. 20 a 172. 16 3. 35 a 1. 511 0. 293 a :, E I EF E I EF,, (Α= 0. 05 D ata are p resented in the fo rm at of m ean standard erro r; sam e letter deno tes non2signifi2 cant difference w hile different letters deno te a significant difference (Α= 0. 05 ( 2 3 T albe2 and T able3 are the sam e as T able 1 ( 2 B E I EF ( 1, E I EF E I EF 1, E I EF,, E I, EF 2. 2, EF E I ;, E I EF ;, 9 00 E I EF ( 3 A ( 2,, E I EF (p < 0. 01 E I EF,, EF E I ( 3 B ;, E I EF ( 2 E I EF 2 EF, E I (p < 0. 05, EF 2. 3, E I ( 4 A E I EF ( 3 : E I,,,, ; EF,, 3 (11 2 F ig. 3 D iurnal changes of net assim ilation rate and transp iration rate of the trial at the late stage of drought
5 : 541 2 (11 Table 2 values of diurnal pho to synthesis, diurnal transp iration and w ater use efficiency of L olium p erenne at the late stage of drought Physio logical characteristics of E I and EF T rial T rial value of W ater use value of diurnal value of W ater use value of diurnal diurnal efficiency diurnal efficiency pho to synthesis pho to synthesis transp iration (mmo lco 2 (mmo lco transp iration 2 (mmo lco 2 (mmo lco 2 (mo m - 2 lh 2O (mo m - 2 mo lh 2O m - 2 lh 2O m - 2 mo lh 2O 385. 52 45. 74 a 92. 87 2. 04 a 4. 134 0. 409 a E I M ild 351. 37 20. 10ab 72. 97 4. 46 b 4. 985 0. 605 a 122. 47 10. 94 c 26. 23 3. 71 c 4. 951 1. 007 a 3 (11 8 Table 3 values of diurnal pho to synthesis, diurnal transp iration and w ater use efficiency of L olium p erenne after rew atering Physio logical characteristics of E I and EF 388. 56 34. 93 ab 74. 84 6. 00 ab 5. 343 0. 954 ab E I M ild 351. 77 31. 87b 71. 63 4. 26 a 4. 965 0. 463 a 231. 56 17. 22 c 60. 58 3. 82 b 3. 853 0. 371 b 477. 14 60. 13 a 98. 71 7. 21 a 4. 841 0. 539 a 380. 48 23. 47 ab 75. 76 6. 78 a 5. 096 0. 489 a EF M ild 308. 14 15. 23b 75. 26 4. 11 b 4. 126 0. 199 a 58. 70 6. 01 d 24. 80 4. 62 c 2. 445 0. 212 b EF M ild 317. 36 28. 22bc 69. 34 5. 03 a 4. 576 0. 272 a 262. 21 18. 45 c 60. 29 5. 89 ab 4. 415 0. 472 ab E I EF,, E I,, EF E I EF ( 4 B, 2. 4 E I EF ; ( 2, E I EF, ( 5 A E I EF, E I EF 4 (11 8 F ig. 4 D iurnal changes of net assim ilation rate and transp iration rate of the trial after rew atering
542 25 5 ( 14 00 F ig. 5 Changes of net assim ilation rate and transp iration rate of the trial during drought (m easured at 14 00 ;,, EF E I EF, E I ; E I, Belesky (1987 EF, EF EF E I, son (1990, CO 2 ( 5 B, EF,, 4 E I 3,, R ichardson (1990,, E I EF (, 1991;, 1990, ( ( ( E I EF,, (Chaves, 1991,,,, M ark s C lay (1996 E I EF (CO 2,,, CO 2 1989 10 E I EF, 7 12,, EF, M ark s C lay E I EF M ark s C lay (1996 35, (1996, M ark s C lay, CO 2,, 35, CO 2 E I EF Belesky (1987, 35, E I CO 2 EF 20% 25%, Be2 lesky ( 1987 ( F estuca ov ina vaṙ EF R ichard2, E I EF, Belesky (1987 Am alric (1999,, E I EF, E I, R avel brachyp hy lla, (1997, R avel, EF E I,, E I EF
5 : 543, E I,,,,,, E I of L olium spp. EF ( gy, 130: 227 238. (C lay, 1990, E I EF L ew is (1997,,,, drough t. A gronom ie, 17: 173 181. Am alric, C., H. Sallanon, F. M onnet & A. H itm i. 1999. Gas ex2 change and ch lo rophyll fluo rescence in sym bio tic and non2sym bi2 o tic ryegrass under w ater. Pho to synthetica, 37: 107 112. Belesky, D. P., O. J. D evine, J. E. Pallas J ṙ &W. C. Stingeṙ 1987. Pho to synthetic activity of tall fescue as influenced by a fungal endophyte. Pho to synthetica, 21: 82 87. Chaves, M. M. 1991. Effects of w ater deficits on carbon assim ila2 tion. Journal of Experim ental Bo tany, 42: 1 16. C lay, K. 1987. Effects of fungal endophytes on the seed and seedling bio logy of L olium p erenne and F estuca arund inacea. O e2 co logia, 73: 358 362. C lay, K. 1990. Fungal endophyte of grasses. A nnual Review of E2 co logy and System atics, 21: 257 297. D e Battista, J. P., J. H. Bouton, C. W. Bacon & M. R. Siegel. 1990. Rhizom e and herbage p roduction of endophyte2removed tall fescue clones and. A gronom y Journal, 82: 651 654. Hoveland, C. S. 1993. Impo rtance and econom ic significance of the A crem onium p lanṫ endophytes to perfo rm ance of anim als and grass A griculture Eco system s and Environm ent, 44: 3 12. L ew is, G. C., C. Ravel, W. N affaa, C. A stier & G. Charm eṫ 1997. O ccurrence of A crem onium endophytes in w ild in European countries and a relationship betw een level of infection and clim ate in F rance. A nnals of A pp lied B io lo2 M arks, S. & K. C lay. 1996. Physio logical responses of F estuca arund inacea to fungal endophyte infection. 133: 727 733. Q i, Q. H. N ew Phyto logist, (, X. W. Sheng ( & S. J iang (. 1990. T he effect on comm unity pho to synthetic rate of A neu role2 p id ium chinense after irrigation and fertilization treatm ents. Bul2 letin of Bo tanical Research (, 10 (3 : 99 105. ( in Ch inese Ravel, C., C. Courty, A. Coudret & G. Charm eṫ 1997. Beneficial effects of N eotyp hod ium lolii on the grow th and w ater status in perennial ryegrass cultivated under nitrogen deficiency o r Read, J. C. & B. J. Camp. 1986. The effect of fungal endophyte A crem onium coenop hialum in tall fescue on anim al perfo rm ance, toxicity, and stand m aintenance. A gronom y Journal, 78: 848 850. R ice, J. S., B. W. P inkerton, W. C. Stinger & D. J. U nder2 sander. 1990. Seed p roduction in tall fescue as affected by fungal endophytes. C rop Science, 30: 1303 1305. R ichardson,m. D., C. W. Bacon & C. S. Hoveland. 1990. The effect of endophyte removal on gas exchange in tall fescue. In: Q uisen2 berry, S. & R. Joo st eds. P roceedings of the international sympo2 sium on A crem onium grass interactions, Baton Rouge. LA : L ouisiana A gricultural Experim ent Station. 189 193. Siegel, M. R. 1993. A crem onium endophytes: our current state of know ledge and future direction fo r search. A griculture Eco sys2 tem s and Environm ent, 44: 301 321. Sun, G. Y. (, Q. Zou (, B. S. Cheng ( & T. W ang (. 1991. Responses of pho to synthetic rate and stom atal conductance to w ater in soybean leaves. Bo tanica Sinica (, 33: 43 49. (in Ch inese A cta : :