27 2 V o l. 27, N o. 2 2001 3 A CTA A GRONOM ICA S IN ICA M ar., 2001 Ξ 3 1 2 1 R. M. V isp eras 2 1 ( 1,, 225009; 2, ), 6 ( g ) AD PG (EC 2. 7. 7. 2 (EC 2. 4. 1. Q 2 (EC 2. 4. 1. 18) 3 : ; AD PG 3 6d; Q 2 ( 3 12d), Q 2 g, : 3 Q 2 ;, ;, AD PG ; ; Q 2 ; Changes in Activ ities of Three Enzym es A ssoc ia ted w ith Starch Syn thesis in R ice Gra in s dur ing Gra in F illing YAN G J ian2chang 1 PEN G Shao2B ing 2 GU Sh i2l iang 1 R. M. V isperas 2 ZHU Q ing2sen 1 ( 1 K ey L ab of C rop Cu ltivation & P hy siology, M inistry of A g ricu ltu re, Y ang z hou U niversity, Y ang z hou, J iang su 225009, Ch ina; 2 International R ice R esearch Institu te, P h ilipp ines) Abstract Six rice cu ltivars ( including 2 japon icagindica hyb rid com b ination s and 1 new p lan t type line ) w ere grow n in w ater cu ltu re in green hou sego r in po ts in screen hou se. T he activity changes in th ree enzym es of AD Pglu soce pyripho spho rylase (EC 2. 7. 7. 2, Starch syn thase (EC 2. 4. 1. 2 and starch b ranch ing enzym e o r Q 2enzym e (EC 2. 4. 1. 18) in bo th of superio r and inferio r grain s and the relation sh ip to grain filling w ere exam ined. T he changes of th ree enzym e activities w ere associated w ith grain filling p rocesṡ T he tim e reach ing a m ax im um of starch syn thase activity w as a little earlier than o r synch ronou s w ith that of grain filling rate. M ax im um AD Pgluco se pyropho spho rylase activity appeared at 3 6d befo re the h ighest grain w eigh ṫ Q 2enzym e activity reached the m ax im um w hen grain w eigh t w as clo se to the h ighesṫ Each enzym e activity, in term s of the m ean of w ho le filling period, the m ean at early filling stage (3 12 days after flow ering), and the m ax im um, w ere sign ifican tly o r very sign ifican tly co rrelated w ith the m ean filling rate, m ax im um filling rate, Ξ (39970424),, : 1999210225, : 2000205201 Received on: 1999210225, A ccep ted on 2000205201
1 58 27 grain w eigh t and fertilized grain2filling percen tage. Q 2enzym e had the largest coefficien ts in the th ree enzym es, on the w ho le. T he enzym e activities in the grain s of japon icagindica hyb rid com b ination s w ere associated w ith their paren tṡ R egu lating sou rce2sink relation sh ip s o r nu trien t level in the p lan t by leaf2cu tting, sp ikelet2th inn ing o r n itrogen app lication at heading cou ld increase o r reduce the enzym e activities in grain s, especially in the inferio r at early filling stage. T he resu lts indicated that the th ree enzym es, especially Q 2enzym e, p lay an im po rtan t ro le in rice grain filling. It is po ssible to breed a variety o r a hybrid w ith good grain2 filling by u sing the paren ts w ith h igh activities of the enzym es in grain s, and to increase grain filling percentage and grain w eigh t by increase of the enzym e activities in grains at early filling stage. Key words AD Pgluco se pyropho spho rylase; Starch syn thase; Q 2enzym e; Grain filling 90% [ 1 ],, [ 2, 3 ] ( ), [ 4, 5 ],, AD PG (AD Pgluco se pyropho spho rylase, EC 2. 7. 7. 2 (Starch syn thase, EC 2. 4. 1. 2 Q 2 ( starch b ranch ing enzym e o r Q 2 [ 6 8 enzym e, EC 2. 4. 1. 18) ] N akam u ra T sai M atters 1 3, Q 2 [ 9 11 AD PG ], 3 ( ),,,, 6 ( g ) AD PG Q 2, 1 1. 1 ( ) IR 72 IR 346862179212221R ( P IR 5729821742 121R ( P2), IR 72083A gir 346862179212221R ( JgIH IR 72083A gir 5729821742121R ( J gih 2), IR 65598211222 (, N PT ) 1. 2 1998 (7 11 ) (14 11 N, 121 15 E), : 7 31 25 ( 0. 28m, 0. 93m, 2. 5m ) 0. 10m [ 12 0. 15m, 1, 50 M ae ], 1 m o lgl HC l 1 m o lgl N aoh ph 5. 0,,, 1g4 : 0. 24m, 0. 245m 0. 16m, M aahas (A ndaquep ti H ap laquo ll) 4 kg N P 2O 5 K 2O 2g 0. 36g 1. 2g, 25, 7 31, 3, 1 20
2 : 3 159 1. 3 5, 1g, 16, 8 1g2, 8 1g2 1. 4 120 150,, 21d 3d 24d 6d, 10 12, ( 1 2d ) ( 2d ), 30s, - 70,, 70, (30 50 ) 1. 5, R ichards [ 13 ] Q 2 AD PG ( ) [ 6 N akam u ra ] [ 14, ], : (% ) = g 1. 0 100%, 2 2. 1 R ichards 1, Cultivars 1 (FGFP) (GW ) Table 1 Fertilized gra in-f illing percen tage (FGFP), gra in we ight (GW ) and some gra in-f illing parameters of test cultivars R 0 Fm ax Po tential (m g d - 1 grain - 1 ) (m g d - 1 grain - 1 ) Fm ean (d) D (d) Tm ax. f (d) FGFP (% ) GW (m g grain - 1 ) IR 72 S 0. 6497 2. 1156 0. 8993 27. 5 6. 85 98. 32 24. 73 I 0. 3526 1. 4268 0. 6922 30. 6 10. 22 87. 41 21. 18 JgIH 1 S 0. 6831 2. 8104 1. 1119 27. 6 9. 38 95. 78 30. 69 I 0. 5568 2. 2562 0. 7643 34. 2 15. 17 86. 43 26. 14 P1 S 0. 9645 2. 6067 1. 4357 21. 3 6. 64 96. 45 30. 58 I 0. 6606 2. 2108 0. 9936 28. 1 13. 50 90. 49 27. 87 JgIH 2 S 1. 3606 2. 9351 1. 3284 21. 1 5. 52 92. 31 28. 03 I 0. 4435 1. 3372 0. 5601 39. 5 21. 06 80. 49 22. 12 P2 S 0. 9846 2. 4631 1. 2531 20. 9 6. 12 93. 59 26. 19 I 0. 5037 1. 5308 0. 5652 36. 2 15. 78 82. 14 20. 46 N PT S 0. 4536 1. 7532 0. 6666 39. 2 13. 25 75. 12 26. 13 I 0. 3495 1. 5534 0. 5416 43. 3 16. 17 69. 43 23. 45 S: Superio r grains; I: Inferio r grains; R 0 : Initial grain filling po tential; Fm ax: M axim um grain filling rate; Fm ean: M ean grain filling rate; D: Grain filling duration; Tm ax f: T he tim e reach ing the m axim um filling rate; FGFP: Fertilized grain filling percentage (fertilized grain w eigh t full filled grain w eigh t 100% )
1 60 27, 6 3 : (R 0 ) (Fm ax) (Fm ean) (D ),, IR 72, IgJH 1 P1; R 0 Fm ax Fm ean D,, Jg IH 2 P2; R 0 Fm ax Fm an,,, N PT 2. 2 Q 2 AD PG,,,, ( 3 12d) 3, (13 26d) (27 39d),, ( IR 72 JgIH 1 P,,,, g J gih 1 J gih 2, P1 3 P2, JgIH 1 J gih 2, 3,, ( 3 12d, 9 18d) ; AD PG, 3 6 ( 9 15d, 12 21d) ; Q 2, ( 12 18d, 15 27d), 1 3, 2. 3 3 2 Q 2 AD PG ( r ( r2) (r3) (r4) 2, ( 3 12d), ( 13 26d) ( 27 39d) Q 2 AD PG 3,, Q 2, AD PG, Q 2 2. 4 3 1g2 1g2, 3,,, ( ),, IR 72 JgIH 2 N PT 3, IR 72,, J gih 2, N PT : IR 72 3 (1. 53sp ikeletsgcm 2 leafarea),
2 : 3 161 1 3 F ig. 1 Changes in activities of three enzym es in the grains during grain filling V ertical bars rep resent standard erro rs (n= 4), ; J gih 2 (1. 67 sp ikeletsgcm 2 leaf area), 1g2,,, 1g2,, ; N PT (1. 61 sp ikeletsgcm 2 leafarea ),,
1 62 27 2 3 (r (r2) (r3) (r4) Table 2 Correlation coeff ic ien ts of the activ ities of the three enzymes in r ice gra in s with the max imum f illing rate (r, mean f illing rate (r2), f illed gra in we ight (r3) and fertilized gra in f illing percen tage (r4) Enzym e activity DA F r1 r2 r3 r4 Q 2 3 12 0. 8513 3 3 0. 9404 3 3 0. 7586 3 3 0. 7890 3 3 Q 2enzym e 13 26-0. 1902-0. 4176-0. 0018-0. 2769 (units m in - 1 grain - 1 ) 27 39-0. 7151 3 3-0. 6920 3 3-0. 5823 3-0. 8059 3 3 M ean 0. 8064 3 3 0. 6915 3 3 0. 8794 3 3 0. 6809 3 3 M axim um. 0. 9242 3 3 0. 8265 3 3 0. 8780 3 3 0. 7755 3 3 3 12 0. 8285 3 3 0. 8573 3 3 0. 7372 3 3 0. 8733 3 3 Starch synthase 13 26-0. 4715-0. 6263 3-0. 2090-0. 5791 3 (nmo l m in - 1 grain - 1 ) 27 39-0. 6329 3-0. 6517 3-0. 5274-0. 5235 M ean 0. 7281 3 3 0. 6938 3 3 0. 7662 3 3 0. 7896 3 3 M axim um 0. 7597 3 3 0. 6217 3 0. 7380 3 3 0. 7486 3 3 AD PG 3 12 0. 6625 3 0. 7552 3 3 0. 5881 3 0. 7863 3 3 AD Pgluco se 23 26-0. 6501 3-0. 6948 3 3-0. 4575-0. 6390 3 Pyropho spho rylase 27 39-0. 7371 3 3-0. 7626 3 3-0. 7351 3 3-0. 7538 3 3 (nmo l m in - 1 grain - 1 ) M ean 0. 5847 3 0. 6885 3 3 0. 6120 3 0. 7574 3 3 M axim um 0. 6905 3 3 0. 7534 3 3 0. 7186 3 3 0. 8292 3 3 DA F: D ays after flow ering; 3 0. 05 Significance at 0. 05 level; 3 3 Significance at 0. 01 level. 0. 01, N PT 1g2, 3 ( 3), N PT,,,, Q 2 AD PG, 3 3 Q 2 AD PG,, ( ), 3 Q 2 AD PG 3 [ 15, 16, AD PG ], [ 17, 18 AD PG, ], 3, Q 2, Q 2 Α21, 6, Α2, AD PG, [ 19 21 ],, Q 2, AD PG,,,
2 : 3 163 3 3 Table 3 Effects of source- sink and n itrogen treatmen ts on the activ ities of three enzymes [SSase: starch syn thase; AD PGP: AD PG AD Pglucose pyrophosphorylase; Q- Q-enzyme. Values Cultivars expressed as un its m in - 1 gra in - 1 (Q-enzyme) and nmol m in - 1 gra in - 1 (SSase and AD PGP) ] T reatm ents Superio r grains Q 2enzym e SSase AD PGP GW (m g) PGFP (% ) IR 72 CK 647. 7 5. 52 67. 17 24. 67 97. 53 1g2 1g2 leaf2cutting 607. 4 4. 87 3 60. 87 23. 47 3 92. 13 3 1g2 1g2 sp ikelet2thinning 654. 7 6. 01 69. 05 25. 26 98. 27 N app lication 650. 0 5. 61 67. 47 24. 93 98. 23 JgIH 2 CK 813. 2 5. 28 55. 73 28. 23 96. 67 1g2 1g2 leaf cutting 760. 1 3 3 3. 83 3 3 48. 53 3 3 26. 11 3 3 92. 91 3 1g2 1g2 sp ikelet2thinning 830. 3 5. 54 55. 80 28. 47 97. 17 N app lication 821. 9 5. 47 59. 33 28. 90 97. 50 N PT CK 303. 7 3. 73 52. 27 25. 70 75. 70 Cultivars 1g2 1g2 leaf2cutting 269. 0 3. 13 3 47. 17 23. 80 3 72. 47 1g2 1g2 sp ikelet2th inning 365. 0 3 3 4. 61 3 3 65. 01 3 3 26. 91 3 81. 36 3 3 N app lication 322. 45 4. 05 54. 27 26. 23 78. 27 T reatm ents Inferio r grains Q 2enzym e SSase AD PGP GW (m g) PGFP (% ) IR 72 CK 420. 5 3. 71 56. 23 21. 40 87. 47 1g2 1g2 leaf2cutting 384. 2 2. 28 3 3 48. 24 3 18. 08 3 3 82. 00 3 3 1g2 1g2 sp ikelet2th inning 464. 9 4. 18 64. 60 3 22. 27 90. 10 3 N app lication 453. 3 4. 06 59. 67 22. 17 90. 37 3 JgIH 2 CK 289. 8 3. 15 12. 66 22. 31 80. 19 1g2 1g2 leaf2cutting 172. 6 3 3 1. 80 3 3 8. 57 3 3 17. 71 3 3 72. 47 3 3 1g2 1g2 sp ikelet2th inning 408. 1 3 3 4. 44 3 3 40. 76 3 3 25. 46 3 3 86. 52 3 3 N app lication 401. 5 3 3 3. 83 3 25. 23 3 3 23. 53 3 84. 03 3 N PT CK 214. 6 1. 53 24. 67 23. 40 67. 83 1g2 1g2 leaf2cutting 134. 1 3 3 0. 90 3 14. 97 3 3 19. 53 3 3 57. 28 3 3 1g2 1g2 sp ikelet2th inning 321. 6 3 3 3. 15 3 3 43. 57 3 3 24. 90 3 3 75. 37 3 3 N app lication 289. 6 3 3 2. 15 3 33. 17 3 3 24. 17 72. 03 3 3 3 6 9d 3 Each value of the activities of th ree enzym es rep resents the m ean of th ree determ inations at 3, 6 and 9 days after flow ering; PGFP: Fertilized grain2filling percentage; GW : Grain w eigh t (m g grain - 1 ) ; 3 V ery significant difference w ith the contro l Significant difference w ith the contro l; 3 3, ( ),,,, (, [ 22 ) ], : Q 2 AD PG ;,
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