An evaluation of lupins (Lupinus spp ) for seed protein production : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy at Massey University, Palmerston North, New Zealand

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(1)Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author..

(2) l.LJI'INS. AN EVALUATTON OF FOR. a. PROTEIN. thesis presented. the of. SEED. requi r ements. in. for. the. North,. spp.). P�ODUCTION. partial. Philosophy at Mas sey. Pa1me rst o n. (l,ur,inus. fulfilment o f. degree o f U n i v e r s i ty,. New Zealand.. NEV1LLE JOHN \.JTTfiERS. Doctor.

(3) A B STRACT. S i nce 1 9 7 2 ther e has b e en s e e d p ro t e i n i n g r ai�b a s ed. i n i t i a t ed. to. l. Lup ins wer e one. r e q u i r emen t .. this. Th i s s t ud y was. p rov i d e in f o rma tion on the agr onomic r equi rement s o f. Lupinus angustifo l ius,. L.. luteus. wi th e mp h a s i s on the s o u t h e r n a d d i t i on ,. f o r s t ock .. meals. fil. o f the c ro p s pro p o s e d t o. in the grea t e r u s e o f. interest. s ome mo r e ba s i c. ca r r i ed out t o p r ov i d e. North. studies. l o c a t i on and the r e s p o n s e of. albus f o r s e e d p r oduc t i on. fs land o f New Zea l and .. In. c a r bon and n i t ro gen t r ans. on. lupins. better. a. L.. and. t o wa t e r s t re s s wer e a l s o. und e r s t a ndi ng. o f t h e l up i n p lan t. an d i t s r e sp o n s e to i t s envi r onme n t .. Ini t i a l l y some fie l d. experiments were. r e s p o n s e s to sow i n g date ,. plant. density,. A t w i d e s pac i ng, L. aYI.(.IUStifol?:us d e crease Oc t o b e r .. in. seed. yield/plant a s. A t no rma l. b e s t s e ed y i e l d . c on c l uded tha t,. densities,. showed. l a id d own t o measure defo l i a t ion a n d c u l t iv a r . a n approxima t e l y l inear. sowing date. h ow ev e r. moved from Ap r i l to. , sowing i n l a t e July gave the. Aut umn sow i n g s we r e a f f ec t ed by d i s ea s e . in. I t wa s. the a b s enc e o f d i s ea s e , seed y i e l d was l a r g e l y. d e t e r mined b y t he l e n g t h o f t he pe r iod o f favo u rab l e env i ronmen t a l c o n d i t i ons be tween the s t a r t o f f l ow e r i rig and t h e f i n i sh o f r e p r o d u c t i v e d eve l opme n t . l a t e ra l in f l oresc en c e s. This. produced. pe r i o d d e t ermined the numb e r o f which , i n tur n , d e t e rmined the n umb er. of pods p r od uc ing seed .. Pod n umbe r was the ma i n component. i n f l uenc ing seed y i e l d .. Th u s , ear l y sowing and r e l ia b l e s umme r. r a i n fa l l or i r r i ga t i on s e em t o b e. the. f a c tors d e t ermining h i gh. l u p i n s e e d y i e l ds .. Respon s e s to den s i t y we r e v a r i a b l e . n o r e sponse in seed y i e ld by. four. i n c re a s e s in seed y ie l d we r e 2 f rom 2 5- 1 00 p l / m .. obtained. In one exp e r iment t h e r e was. c u l t ivars over th e s e sow i n g t ime s 2 to d en s i t i e s rang i n g f r om 50- 1 4 0 pl / m . In a f ur th e r exper imen t , a s p l a n t d e n s i ty i n c r eas e d.

(4) Remova l o f the main s t em g r owi ng p oi n t ear ly i n growth b r i e f l y s t imu l a t e d l a t e ra l s t em grow t h b u t t h e e f f e c t on l a t e r a l s tem seed y i e ld wa s i n suf f i c i e n t. c ompen s a t e f o r the loss o f the main. to. s t em seed s .. The re was l i t t l e. difference. Uni h a rve s t , U n i whi t e a nd s pr i n g s ow i n g , Un i c rop under d ry. t h e L. angustifo lius c u l t iv a r s. be t ween. Unicrop. flowered. wh e n s own e a r ly but , wi t h l a t e e a r l i e r wh i ch w a s a n advant age I n one exp e rime n t c omp a r ing. e a r l y summe r c o ndit i on s .. a r ange o f l e g ume s p e c ies , L. albus and Pisum sativum p r od u c e d t he. h ighes t s e ed y i e l d b u t L. albus a n d L. p ro t e i n p e r uni t a r e a .. The. peak. l uteus y i e l d e d t h e mo s t. r a t e o f nit r ogen a c c umu l a t i on. i n a l l sp e c i e s wa s s i mi l a r a nd t h e. main. f a c t o r i n f l uen c i n g p ro t e i n. y i e ld a p p e a r e d t o b e t h e dura t ion o f n i t rogen ac cumul a t i on . Provided e a c h c rop u t i l i s e d s i m i l a r d u ra t i o n s of th e growing p e r i od , t he yie l d o f s e e d p r ot e i n / ha f rom va r i ous l e gume c r op s i s l i k e ly t o be s i m i l a r ; seed .. t h e ma i n d i f f e r e n c e be i n g. I t wa s sugge s t e d. the. compos i t i o n o f the. f o r max i mu� seed p r o t e i n yi e l d,. that,. i n de t e rm i n a t e c u l t i va r s ma y have s ome advan tage over more d e t e rmi na t e c u l t iva r s p rov ided app r op r i a t e mana gement p r ocedures a r e ado p t ed .. S t ud i e s on wa t e r s t re s s indica t ed th a t b y i n f l ue n c i n g t h e d i s t r i b u t i on r e p rodu c t i ve g row th.. Mild. of. wa t e r. g r owth and i n c r e a s e t h e r a t e. it. pl ays an i mp o r t a n t r o l e. assimi l a t e be tween vege t a t ive and. stress. o f seed. t ended t o s t op vege t a t ive. g r owt h .. !fuen. s u f f i c i en t l y. seve r e , wa t e r s t r e s s appeared t o i ni t i a t e the s e n e s c e nc e o f t h e p lan t , t h e t i m i n g o f wh i ch t h a t s i t ua t i on .. Wa t e r. reduc i ng p od number,. determined. deficit. t he p o t e n t i a l s e e d y ie l d f o r. h a d i t s ma i n e f f e c t o n s e e d y i e l d by. Othe r y ie l d compone n t s wer e re l a t i v e l y s t able.. Day t emp�rat ures o f 28°C, whe n imposed ea r l y i n g r owt h , r e d uce d. vege t a tive a nd s eed y i e l d i n L. albus.. As the pla n t d eveloped,. h oweve r , the a dver se e f f e c t s of h i gh temperature d e cre ased un t i l.

(5) g rowth wa s s t imu l a t ed. d11ring. f i r st o r d e r l a t e r a l f l ower i n g .. No d i rec t e f f ec t o f h i gh temp era t ur e on pod a b s c i s s i on was a p p a r en t and i t wa s s ug g e s ted t h a t pod l o s s und e r h i gh t emp e r a t u r e s whi ch have been r ep o r t ed o c c ur red l a r ge l y. because of. an a s s o c i a t ed. wa t e r s t r e s s . A. 14. C t ra n s l o c a t i o n s tu d y i n dica t ed t h a t mo s t movement of. pho tosyn t h a t e in L. albus was in t o the b ranch on wh i c h the l ab e l l ed l e a f was i n s e r t ed , o r i n t o to i t .. lower. b ra n c h o r d e r s di r e c t ly c onnec t ed. R e s u l t s sugge s t t h a t , in L. albus c v . U l t ra , lower or d e r. s t ems a r e a mor e i mp o r t a n t c ompet i t or w i t h t h e i n f l o r e s c en c e f o r p h o t o sy n th a t e t han t h e new ,. ra p idly d ev e lop i ng, h i gh e r o r d e r l a t er a l. b ra n c h e s . A p o s s ible stra t egy for growing l u p i n i n a comme r c i a l ly v i ab l e. s i tua t i o n. in. t he S o u t h e rn North I s l an d i s d i scussed ..

(6) TABLE OF CONTENTS PAGE 1. I NT RODUCTI ON : S ECT I ON A :. LITERAT1JRE REVIEW. 5. A. 1. See d Produc t i on in Annua l Le gumes. 5. A.2. Bac kground to Lup i n G rowing. 6. A.2 . 1. Lup in Seed Qua l i t y. 7. A.2 .2. Lupin S p e c i e s. 8. A.2.2. 1. Lupinus angus t i f o l i us. 8. A. 2 .2 . 2. Lup inus a lbus (wh i te l up i n ). 9. A. 2 .2 . 3. Lup i n u s l u t eus (ye l l ow l up i n ). A. 3. A.4. The Deve lopme n t o f Legume Seed Y i e ld. 11. A . 3 . 1 P l an t S t r uc t ur e a n d Developmen t. 11. A . 3 . 2 Compone n t s o f L e g ume Seed Yie l d. 14. A . 3 .3 F l owe ring and Pods e t. 16. A . 3 . 4 Pod a n d Seed Deve l opme n t. 19. The Carbon and Ni t r o ge n Economy o f G r a i n Le g umes. 20. A . 4 . 1 The Carbon Ec on omy. 21. A . 4 . 2 The Ni t rogen E c on omy. 23. A.4 . 3 A.5. 10. A. 4 . 2 . 1. I n i t i a l As s imi l a t ion. 24. A. 4 . 2 . 2. U t i l i s a t i on o f Ni t rogen. 26. The Imp o r t ance of Sene s ce n c e in Legume Seed P rodu c t i on. 27. S ome Ef f e c t s o f Envi ronment on S e e d Y i e ld. 32. A. 5 . 1. 32. Wa t e r D e f i c i t and Y i e l d A.5 . 1 . 1 A.5. 1.2. A . 5. 2. The Main Phy s i o l o g i c a l E f f e c t s o f Drough t. 33. Ef fe c t o f Wat e r S t r e s s o n Legume Seed Prod uc t ion. 35. Re la t i o nship be tween Temperat u r e and See d Y i e ld. 37. A,5.3. Time o f Sowing. 39. A. 5 . 4. P l a n t Den s i t y E f fe c t s on S e e d Y i e ld. 41.

(7) PAGE S ECT I ON B : B. 1. B.2. B.3. B.4. FXELD TRIALS. In f l uence o f T i me o f Sowing on See d Yi e l d. 44. B. 1 . 1. I n t roduc t i on. 44. B. 1 .2. Ma t e r i a l s and Me thods. 44. B. 1 .3. Re s u l t s. 46. B.1.4. D i s c us s i o n. 51. Th e E ffe c t o f Sowi n g T i me and P lant Pop u l a t i o n on Fo u r Cu l t i vars of Lup i ns. 52. B.2 . 1. I n t roduc t i on. 52. B.2.2. Ma t e r i a ls and Me t h o d s. 53. B.2.3. Re s u l t s. 54. B.2 . 4. D i s c us s i on. 56. A Spac i n g and D e f o l i a t i on S t udy. 59. B. 3.l. I n t rod uc t i on. 59. B.3 . 2. Ma t e r i a l s and Me t h o d s. 60. B . 3.3. Re s u l t s. 61. B.3 . 4. D i s c us s i on. 71. A Compa r i s on o f s e v e r a l g r a i n legume s at t wo sowing t i mes. 73. B.4 . 1. Seed Y i e l d and C omponen t s. 73. B . 4 .1 . 1. I n t r o d uc t i on. 73. B.4 . 1 .2. Ma t e r i a�s and Me t h o d s. 75. B.4 . 1 .3. Res u l t s. 76. B.4 . 1 .4. D i s c us s i on. B. 4 . 2. D i s t r i b u t ion o f N i t rogen w i t h i n ab ove- ground c omp onen t s. 83. B . 4 .2 . l. I n t ro d u c t ion. 83. B . 4 .2 . 2. Ma t e r i a l s and Me t h o d s. 83. B . 4 .2 .3. Re s u l t s. 85. B.4 . 2 .4. D i s c us s i on. 103.

(8) PAGE. S ECTION C : C.1 C.2. WAT ER STRE S S STUDIES ON LUPINUS ALBUS Gene r a l I n t ro d uc t i on. 108. E f f e c t o f wa t er s t ress i mp os e d dur i ng a s i n g l e growth s t age a t two h umi d i ty leve ls. 1 10. C . 2.1. I n t roduc t i on. 1 10. C.2.2. Ma t e r i a l s and Me t h o d s. 111. C.2.3. Resul ts. 1 15. C.2 . 3 . 1. C , 2,4 C, 3. of. growth s t ages. 115. C.2 . 3 . 2. Dura t i on. C . 2 .3 , 3. To t a l p l an t grow t h. 117. C.2,3.4. Lea f growth. 1 19. C . 2 .3 . 5. S t em G row t h. 123. C.2. 3.6. Reproduc t i ve growt h. 1 25. D i s c us s i on. 1 15. 131. E f f e c t of w a t e r s t r es s imposed d u r i n g two o r th ree growth s t age s. 1 36. C . 3.1. I n t r oduc t i on. 1 36. C.3.2. Ma t e r i a ls and Me t h o d s. 1 37. C.3.3. Re s u l t s. 1 40. C. 3.3. 1. Vege t a t i ve growt h. 1 40. C. 3. 3.2. F lowe r n umber. 143. C. 3 . 3. 3. Seed y ie ld and c omponen t s. 144. C . 3.3.4. E f f e c t o f wa t e r s t r e s s on rep roduc t ive gr owth. 1 48. S o u r c e o f n i t r o gen for s e e d growth. 151. C. 3.3.5 C.3.4. C.4. Leve ls o f w a t er s t re s s i mposed. D i s c u s s ion. 1 54. C. 3.4.1. Seed Yie l d. 154. C.3.4.2. E f f e c t o f wa t er s t re s s on as s imi l ate d i s tribution. 156. E f fe c t o f h i gh t empe rature und e r c ondi t i ons 159 o f adeq u a t e and re s t r i c t ed wate r s up p ly C.4 . 1. I n t ro d uc t i on. 15 9. C. 4 . 2. Ma t er i a l s and Me t hods. 160. C. 4 . 3. Results. 161. C.4.4. D i sc u s s i on. 166.

(9) PAGE. SECT I ON D : D.l. S E CT I ON E :. TRANSLOCATION. STLJDY. T r an s l oca t ion. of. 169. 14c. in. Lupinus a lbus. 169. D.l.l. I n t ro d u c t i on. 169. 0. 1 . 2. Ma t er i a l s and Me t h o d s. 170. 0. 1 . 3. Resu l t s. 172. 0. 1 . 4. D i s c us s i o n. 181. C ONCLUDING DI S CU S SION. 185. E.l. Bas i c c o nsi d e r a t i ons. 1 85. E.2. S ow i n g t i me. 1 89. E.3. Lu p i n s p eci e s. 191. E.4. Plant dens i t y. 192. E.5. P o t en t i a l f o r. R EF ERENCES. Lupin. in. t h e Nor t h I s land. 193. 196.

(10) ACKNOWLEDGEMENTS. F i nanc i a l s u p po r t f ro m r e s e a r c h c o n t ra c t s wi th D a l g e t y ( N . Z . ) L t d , a nd D S I R and f rom gran t s b y the PaJme r s to n No r t h C i ty Corpora t i on E c o n omi c Dev e l o pmen t and t h e Pork Indus t ry Council t o o b t a i n mo r e in f o rma t i on t h i s a s s i s t anc e i s. itl. ma d e i t p o s s i b l e. t h i s, and o th e r s t ud i e s on l u p i n s and. gra t e fu l l y ackno wledged . lL R .. To my s up e rv i s o r s , Pr o f .. wi l l ingly p ro v i ded gu i d an c e. and. \.Ja tkin and D r sup po r t ,. I. B.J.. Forde who so. wou l d l i ke to exp r e s s. m y g r a t e f u l t h ank s .. Tech n i c a l a s s i s t an c e wa s p r ov i d ed a t c r i t i ca l t imes b y a n umb e r o f p eo p l e bu t in pa r t i cul a r. I. wi sh t o thank Lo i s Ma t h e r and Terry Lync h. who p rov i d ed re l i a b.le a nd va l uab l e a s s i s ta n c e ov e r mu c h o f the e x p e r imen ta l pe r i o d a n d to. I. am. gra t eful. fo r. Joan. Hay. f o r t yp in g th e t h e s i s ,. t h e u s e of c on t ro l l ed c l i ma t e f a c i l i t i es. at the P l ant Phy s i o1o gy Div i s i on , DS IR and I wou l d l i ke to thank Mrs L i z Edge , Mr Ian Wa r r i n g t o n and other memb e r s of the C l ima t e Ro om S up po r t G roup f o r t h e i r h e l p and in t er e s t .. F ina l ly a b i g t ha nk you to my wife, Ma r i e , and my three boys for the i r pa t i e n c e and sup po r t n e g l e c t i ng them fo r so l ong .. .. My hea r t f e l t a p o l o g i e s a l so f o r.

(11) LI S T OF TABLES. TAB L E. B.1.1. PAGE. S ow i n g and. flowe r i ng da t es,. a n d deg ree-da y s. t o f lowe r i ng. day s. t o f lower i n g f o r Un i c rop and 45. Un i harve s t. B. l .2. Res u l t s o f l i n e ar regre s s i o n ana l y s i s o f s e e d we i g h t and. B.2. 1. t im e. pe r p l a n t. from. S e e d weig h t. B. 2.2. Seed. pe r. m. B.3. 1. num b e r per p l a n t 49. 2. a nd per p l a nt; p roduced s e e d. p e r c e n t age for t h re e 55. da t e s.. we i gh t. pe r. m. of sown p l an t s '�h c ul t. pod. sowing.. o f s own p l a n t s wh i c h s owing. ,. i v a r at. Ef fec t of o f l a t era l. 2. Lch. each. p l a nt. and pe r p l a n t ; pr(ldUI'�d. sowing.. spaclng. seed. on the d r y. ste m s pe r pla nt. f r om. p e r c e n t age for. Pach. 55. we i gh t. e ar l y 61. harves t s .. B. 3.2. B. 3.3. E f fec t. of. spacing on. s t ems p e r. plan t. E f fec t o f. spa c i ng on. i n f l ore s c en c es. t h e numb er of l a t e r a l. f rom e a rly. per. t he. harves t s. number o f f e r t i l e. p l a n t f o r each bran ch. order - f i n a l harves t .. B.3.4. 62. 63. E f f e c t o f s p ac i n g on y i e l d per p l ant a nd c omp onen t s. of. y i e l d - f i n a l ha rves t. 64.

(12) TABLE. B. 3 5 .. PAGE 2 Y ie l d o f ove n d r ied s e e d ( g /m ) - f i n a l h a r ves t. B. 3 . 6. 65. Seed y i e l d ( g ) f or e a c h branch o rd e r a t each s p aci ng and sowi n g , and numb e r o f p od s on mai n s t em and f ir s t ord e r l a t e r a ls o f t he unde f o lia t ed a u t umn-s own p l an t s on 1 8 Oc t o b e r .. B. 3 . 7. 66. E f f e c t o f d e f o l i ation t r e a t me n t s on t he n umbe r o f l a t era l s t ems per p la n t - e a r ly ha rves t s .. B. 3 8 .. 67. E f f e c t of t h e d e f o l i a t ion t rea tmen t s on t h e d ry we i g h t. ( g ) o f l a t e ra l s t ems p e r p l a n t. - e a r l y harves t s. B.3.9. 68. E f fec t o f t h e d e f o l ia t Lon t r ea t men t s on t he 2 t o t a l lea f a r e a p e r p l a nt ( cm ) and l e a f a re a of l a t e ra l s t em s - e a r l y harve s t s. 8 . 3.10. 69. E f f e c t o f t he d e f o l i a t i on trea tment s on t he numbe r o f f l ower site s and p od s f or e ach b ranch o r d e r - e a r l y h a r ve s t s. B.3.11. E f f e c t o f d e f o l i a t ion o n yield p e r p lan t and 71. yie ld compone n t s. 8 , 4. 1. B.. 4.2. B.4. 3. 70. 2 Tot a l s eed y i eld ( g /m ) and t he numb e r of plant s con t ribu t i ng. to. y i e ld. 77. Number o f pods per p l a n t. 78. Numb e r o f s e e d s p e r p od. 78.

(13) TABLE. PAGE. B.4.4.. H un dred seed. B. 4 . 5. N i trogen. B.4 . 6. Y ield. B.4 . 7. Pa t h. B. 4 . 8. Re l a t i ve. (N. o f p ro t e i n. samp l e d 8 - 9 Regr e s s i on eq ua t ions. 2. ). 80. c omp onen t s on y i e ld. content of. water. ( g /m. 6. 2 5 ). x. c oeff i c ien t s of. n i t rogen. 79. per centage o f seed. s amp l e d 1 2 - 2 p � .. B.4.9. 79. (g). weight. upper. c anopy l ea v e s. i n pa re n t he s i s. Results. 88. am.. coefficient. b for regr e s s ion. total nitrogen d u ring. and. non - seed. decline phase \vith. the. d ays 92. from s owi n g .. B.4, 10. Ra t i o of non-see d n it r o g e n we i gh t. (mg / g ). at. the. to. non-seed d r y. f i na l s amp l i n g o f e a c h 94. c u l t i var. B . 4,1 1. Regres s i on. coefficient. b. equa t ions l ea f a n d stem f r om sowi ng a nd. c oe. for reg r e s s i on nitrogen with. d ays. f f i c i e n t c f or q u ad r a t i c. l og i s t i c eq ua t i o n s s t em n i t rogen. with. days. f r om l a t e s ow i n g .. B 4 12 .. .. 80. 96. Regre s s i on coef f i c i en t. b. for regre s s i on. e q ua t i on s pod n i t rog e n a n d seed nit r ogen 100. w i th d ays f rom sow i ng .. B.4 . 13. Es t i ma t ed from p l a n t. contriburion parts and. from a s s i m i l a t i on. of n i t rogen t o s e e d. t h a t sup p l i e d d i re c t 1 02.

(14) TABLE. PAGE of. C.2 . 1. Sched u l e. C . 2 .2. Schedu l e o f harve s t s. C.2.3. Dry weigh t o f ma i n s t em and f i r s t. t r ea t me n t s. 1 13. 114. order l e a f and dry weigh t p e r l e a f a t t h e end o f f i r s t o rde r la t e r a l f l ower ing .. C. 2 . 4. 1 21. Dry we i gh t , n i t r ogen p e r c en t age and ni t r ogen y i e l d o f ma in s t e m l e a f and s t em a t t wo h um i d i t y l eve l s. 1 22. C. 2 . 5. Numb er o f s e eds per ma i n s t em pod. 127. C. 2.6. Numbe r o f p r od uc t i v e f i rs t ord e r l a t e ra l s t ems and pods per p r oduc t ive s t em a t f i na l ha rve s t. C . 2 ,7. We i gh t o f ma i n stem s eed per p l ant a t. harve s t 4 ( end o f 1 s t o rde r lat e r a l f l owe r ing ) and a t f i n a l harve s t. C.2.8. 1 28. N umber o f i n c omp l e t e ly f i l led s ee d i n f i r s t o rd e r la t e r a l pods - f i n a l harves t. C.2.9. 1 28. 1 29. P e r c e n t ag e o f n i t r og e n i n s eed - f i na l ha rve s t. C ,2, 10. Numb e r o f f l owe r s per p lant. 1 30. C, 3. 1. Schedu l e o f t r eatme n t s. 1 38. C.3,2. Schedu l e o f harve s t s. 1 39.

(15) PAGE. TABLE. C . 3 .3. Total pl a nt wei.ght. 141. C.3 . 4. T o t a l plant n i t r ogen c o nten t. 141. C. 3.5. Re l a t i ve g r owt h ra t e s d ur i ng 3 f l owe r i ng periods f o r t o t a l p l an t we i g h t and f or. 1 42. vege t a t ive part s o nly.. C . 3. 6. Number o f f l o we r s i tes per p l an t. C . 3.7. N umber. of. main. s t em pods a t f i na l. h a rv e s t a n d fi rst o r d e r l a t e r a l p o d s at harve s t 4 a n d f i na l harves t. C. 3 . 8. Num b e r of seed s seed weight at. C , 3 ,9. Estimated. 142. pod and hundred. per. final. increase. 1 46. or. harvest. 1 47. d ec r e ase o f. n i t r ogen i n pla nt p a r t s ,. loss o f n i t r og e n. i n senescing leaf a n d t otal ne t n i t r ogen accumula t i o n rat e d u r i ng second order. l a t e ra l f l ower i ng.. C . 3 . 10. 1 52. Es t i ma t e d t r anslocation o f n i t r o gen b e tween p l a n t par t s f o r t he p e r i od b e t we e n h a rve s t 4 a n d f i na l ha rves t. C.4. 1. 153. Dry we i gh t o f the t o t a l p l an t , veg e t a t i ve and r e p r od u c t i ve t i ss ue s a t t h e end o f ma i n s t em f l owe r i ng a n d end o f 1 s t o r d e r. C.4.2. la t e ra l f l owe r i ng .. 1 62. Mean d u r a t i o n o f growth p e r i od s. 162.

(16) TABLE. C.4.3. PAGE. Re la t i ve g r owth ra t e s dur i n g 1 s t order l a t e r a l f l ower i ng. 1 63. C.4.4. Number o f f l ower s. 163. C. 4 . 5. We i g h t o f seed p e r p l an t. 164. C. 4 . 6. Numbe r o f pods per p l an t a t f in a l ha rves t .. 164. C.4.7. Numbe r o f s e eds p e r pod a t f i nal harve s t. 165. D. 1 . 1. To t a l a c t i v i t y i n p l an t c omponent s f r om Exp e rime n t 1. D. l.2. Tot a l a c t i v i t y i n p l a n t c ompone nt s f r om Expe r i me n t 2. D .I. 3. 1 75. Tot a l a ct i vi t y i n p l ant c ompone n t s f r om Exp er ime n t 3. D . 1.4. 1 73. 177. To t a l act i v i t y in p l a nt c omponents 2 days a f t e r labe l l i ng f or Expe r i�e n t s 4 - 6 and t he prop o r t ion o f t o t a l a c t i vi ty in each c omponent f or Exp e r imen t s 4 - 7 ,. 1 78.

(17) Ll S T OF. FIGURES. FIGURE. A. 3. 1. PAGE Schematic. layout of. <l. lup i n p lan t show i n g. the p o s i t i o n and n umbe r ing o f t he. branch. orders.. A. 3 . 2. 13. Diagramatic representation o f direct p a t h s. o f i n f l ue n ce of several structural c omp onen t s o f. A. 4. 1. B.. I. .. l. p l a n t u p on yield.. 14. Ou t line o f. the model g iv en b y S in c l a i r. a nd de Wit. (JQ76). Per i od ,. for. da t e , when. B . 1. 2. the. A c c umu l ated of. each. 29. in florescence. �0% o f. >. net. f r os t s per. plants. we re. and s ow i n g f lower ing. 47. w a t e r a va i l a b i l i t y, n umber. week,. and mean weekly air. temperatures for the. 1973/4 sea s o n. 48. B. l . 3. Seed yield p er plant. f or. 50. B, 2 . 1. Total yield and yield o f b r a nch orders. B. 4. 1. Acc um u l a t ed. each bra nch o r d e r. water availability,. 57. and. mean weekly a i r temperature for t he 19 7 4 / 5. 86. s ea s on.. B . 4,2. Dry weight per p l a nr o f and late s o wi n g s. c. om pone n t s. for e a r l y. 89.

(18) FIGURE. B.4 . 3. PAGE. To t a l ni t rogen p e r p l ant and n on-se ed ni t r o g e n p e r p l a n t dur ing t h e de c l ine p h a s e f o r e a r l y and l a t e sowing. B.4.4. N i t r ogen c on t e n t o f l ea f , s t em, pod and s e e d p e r p l an t f o r ear l y -s own p l an t s. B.4.5. Each. p oin t i s a mean o f 8. p l an t s. .. .. 118. To t a l p l ant n i t rogen and p r opor t i on wh i ch wa s s e ed. C 2 4. 1 16. To t a l p l an t d r y weigh t and p r opor t i on wh i ch wa s seed. C.2.3. 99. R e s u l t s o f r e l a t i ve l e a f wa ter c o n t e n t (RWC ) m e a s ur emen t s .. C.2,2. 97. N i t r ogen c on t e n t o f lea f, s t em, p ods and s e ed p e r p l a nt f o r lat e s ow i n g. C.2.1. 91. 1 18. R a t e of c h ange i n l ea f and s t em dry we i gh t and ni t rogen f o r t h e ma i n s t em over three g r owth s t ag e s and f or the f i r s t order l a t er a l s t ems over one g r ow t h s t age. 1 20. C. 2 . 5. Dry weight o f s t ems a t f i n a l harve s t. 1 24. C.2,6. We i gh t of s e ed and numbe r o f p ods p e r p lant. 1 26. C,3, 1. Seed we i gh t p e r p l a nt a t H3, H4 and at f i nal. harve s t. 14 5.

(19) F I GURE. C. 3 . 2. PAGE. Changes harve s t s. C.3.3. in. dry. wei g h t. for control. Cha nges i n d ry. we ig h t. o f c ompone n t s b e t ween p l an t s. o f c omp onen ts b e t ween. h a r v.·�;t s for t r ea t m e n t P F l C.3.4. Ch a n g e s i n d ry. \ve i g ht. of. Changes. in dry weight. 149. c om p o nen t s be t ween. h a r ve s t s fo r t r e a t men t F l 3. C. 3.5. 1 49. 1 50. o f c omponents be t ween. h a r v es t s for t r eatme n t F2'�. 150.

(20) 1. INTRODUCTION. legu mes. Annual grain. as they produce over annually. useage.. However,. th e y. tP x t ,. important as they supp l annually. 1977).. (Sinha,. In New Zealand, cheap. has not. resulted. of the world's protein. onlv Rhnut. a re. 0.75 mi ll ion tonnes of seed. abundant s upp l y of relatively. plant protein,. other than that from pastures,. However,. with a. world-wide shortage. Zealand stork f ee d meals, increased rapidly.. in an awareness that. which were. crops were considered but sweet. known tn he a gond protein source. 1970), became one of the mRin crops c-onsidere d. ,q. Some of the. (a). Are:. ust'. lttpins. �. RS. 1979).. only be grown in the warmer (c). Ability to yield. (d). Better ab l e. intensive cropp i n g. 1970;. Hudson. Gladstones,. subs ti tu te. for. soybean (Williams,. zones of the country.. we]] on. so i l s not norma] l y. (\.laostones. 1970).. suited for. tn utilise phosphate reserves in the soil. compared with many p l an ts. (e). ((aadstones,. 1976) compared 1vi th most grain legumes which provides the. opportunity to. inputs.. (Gladstones,. dv an t ;'l g e s that hAve been noted for lupins as a. High pr o te i n rontent in t he seed. et al.,. This. seed protein may have a place in. A number of. feed mea]s.. seed crop. not very. 1971-72 t he price of meat meals, the main source of. protein in New. lupins,. y. on ly 1R%. lupins (Dur1:n11:� spp.). been very important.. of protein in. stock. supply. because of our. animal protein,. source of protein. 100 million tonnes of hi gh protein food. c on. In this. are an important world. (I.Jhite,. 1961;. Glarlstones et al.,. 1964;. 1970, 1972) p os s i b l y resulting in lower fertiliser. Limited evidence that. fixing nitrogen. (NutmRn,. lupins are one of the better crops at. 197fi;. Boundy,. 1978)..

(21) 2.. Altho ugh lupins were n u t been p reviousl y grown mal n] y (Allen,. 1949).. n. tww. ('rop. a. f<Jrage. as. to. or. and. 1ittl�·. the potential as a s t o ck f o od .. umsider:Jtion had heen given to In. about 170 h a of lupin. 1936-7,. 1 93!)) <lnd th is increased to. was grown for seed (Anon,. about 3 0 00 ha i n 1 9 4 5 ( Whi t e ,. Suhsequen tly,. 1 961).. in the ea r l y 1970's it. wa. s a minor. I nforma t i o n o n lup i n Those references wh i c h. seed. are. prouu('tion. in New. (Anon,. 1956;. \·Jhatman,. and. it. based on farmer experience and. is. the use of. Zealand was limited.. 1938;. McPherson,. 1959;. Hhite,. 1 940 ;. 1 96 1 ). Little basis for their. provided only broad recommendations. recommendations was given. peak of. crop.. a va i labl e. van Steveninck,. a. fe rt i l i t y improved until. lupin for forage declined steadily as soil. 1947 ;. green manure crop. Seed p r o duc tion h :Jd hL"en limited to supplying seed. f or r e so w i n g p u r p oses. Inch,. Z e a l a nd , t h e y had. New. probabl"". dL."signed. for. t h at they were largely Ca n te r b u r y conditions.. Since the main market for l u p in seed destined as stock rations. was likely to be in the No r th Is l a nd , this s t ud y investigated the potential for growing the crop i n t he order to minimise transport costs.. so. uthe rn Nor th I sland in. A l t hough lupin seed production. had been investigated at Mass e y Un i ve r sit y previously (G.S. Robinson,. pers, comm.), no records of tha t w o r k exist. lupins have been grown for. seed. Bitter and sweet blue. on the Manawatu/Rangitikei coastal. sand country mainly as an autumn-sown c ro p often in association with wind erosion control (van Kraayenord, unpublished report).. Because of interest by fanners and feed manufacturers, it was decided to commence a research programme into lupin seed production and after an initial experiment i n. 1 9 7 2 (Withers,. 1 9 7 3 ) the. programme reported in this thesis commenced in 1 9 7 3 .. The overall objective of t he progranune was to provide data which would be useful in assessing the type of environment and management suitable for J up i n seed production that could be u sed as a basis of recommendations to f a rme rs:. and. to. provide basic.

(22) 3. inform a t i on. on. the. g1·m..ttlt. c om p ar i sons wi t h othe r. grain legum�='�,. p r og r a mm e was diviueJ intu. The A. (a). and deve]OfllliL'Ill. se ries. agronomic. field. of simple. \vhicll. trials dL·sign,-d. exper i men t s aimed Lo: compare. c ultivars. t he. u n d er Manaw a t u. s t udies of. ,11'. cha,-ac t e r i s tics. speci e s ,. This. conditions.. so. t hat. the. a11d. r<111gL'. cJ. t he ni t rog e n and d r y 1veight. g rai n l e gume. \vider inte't'est.. h',)tl1d be of. which wotl!d ,·ompl imL·nt. i nforma t ion. i nc lu d i n g. lupins,. ttvo m<�in sections:. conduct e d by o t her 1v11rke rs in An::.l rdl ia. (i). ,,f. p rovide. to \vork. NlJW of. h a sic. was. that. Zealand . lupin. These. reLH ivt:'. rotent i a l. and. species. wo rk included. dist1 ihution. being. of. comp a r a t i ve. lup i ns and orher. of. lupi n s could. e s t ab l i s hed , (ii ). d e t ermine the mus t sown i n autumn. been an. but. the. snuthvrn. area whe r e c ro p s a re sp ring-sown. wh e t h e r ( iii ). lupins. cou l d. i nd ic ate. (iv). (b). yield \vell if. ob t ai n inf o rmat i on. may also requi re. As a. a. tHl. change in. result. of the. l e ngth. of the. have. an. and. (ii). In. occur in late. t r ansl oc a tion of the life of t he. hence. spring. (i),. Early. nit r o geo. fac t or. The. yield.. through. and. wa t e r e d. poss i b J. This. y. were decided. of. dete rmining t he Mana\.;ratu tends. pe r i ods. m ild. str ess. on. was. grow t h. t he ni t rogen. and wa t e r s tres s ed plants. i n d icat e d ea. of. t h e summe r , i t. furr l w r inf ormation on. obse rva t ions. and. t op i cs. envi ronmen t s :. an importan t. rbon. t ha t mas s i ve. \vas occurring during. p lant,. d e nse canopies it w a s noted. s e nesced and canopy.. main. yield should b e de t e rmined.. was to be s t udi ed ,. In. three. densi. ty,. s t ud y ,. t he effects of such short periods. conjun c t ion wi th. t o de t e rm i n e. Changes i.n SOlving tim e. density,. controlled. t o be. t radit iona l ly. impor t ant. Wi1s. usua l l y. spring-sown,. trials,. dist ribu t i on p � t t er n of adeq u ately. (iii). it. erra t i c r a i nfall pattern and as b rief. t ha t. see d. so. Lupins have. l�land i s. more detailed. growi ng season and. water stress can decided. Nnnh. recnmmendL·d plant. fie l,l. Ha ter stress se�med. (i). plant. topi c s worthy of. on for furt her stu d y i n more. to. sowing time.. s uitable. l ow e r. p o d s dtv�lop�d. raised. the question. that. in shade of. l ower lea v es due. whe t h e r. to. t he. often. t he d en s e lower. upper. pods we re. be.

(23) 4.. s up p l i e d f ro m. remobi1Lsed products. c a n o py .. pe ri me n t. An. ex. t r an s l o c a t i on p a t ter n s. T h e M a n a w atu. and. was in. oth�r. in this r eg i o n and , i n f o rma t ion. to. if. l,.. nU•us. low LHHI. e n abl e. whether. so,. directly. therlof,Jre desjgned. I s l an d h a s considerab.le potential was de s i gned t o eva l ua t e. or. a number. ar,�as. fur. <lf. to. t ile. crupping.. ltipiilS. provide. f a r me r s. at. Erow t h e u p p er. sume. i nd i c a t e t h e. of. g r ow th stages.. Soulhern No r th. T hi s p r og ramme. could h e a s u i tabl e c rop of. the. b a sic ag ron omic. t<) Stlt·c·essful'ly grow the c r o p ..

(24) 5.. LITERATURE. SECTION A. RI�V I EW. (I). This review has two ohjectives.. way , t h e maj o r fac tors influendng. examine,. kn o w l e d ge s h o u l d h e l p. in. To rev i ew e xi s t i n g. i t as f a r a s pos s i b l e. to. the. un d e r s t anding l u p in. 011. knowledge. in a gene ral. p r o d u ct i on i n grain. As t h ese crops h ave many f e a t ur e s. l eg ume s .. (2). seed. To. in. common,. of lup i n. s uc h. seed. production .. s e e d product ion an d re l a t e. ge n er a l p r i n c ip l e s o f g ra in l eg ume. the. s e e d p r o d uct i on . Because. of. the wid e. range. of. topics. thi s review is exte ns i ve r a t h er. than. A.1.. LEGUt-1ES. SEED PRODUCTION. IN. ANNUAL. Th e re are maj o r problems wLth. c rop s .. cove r e d i n t h i s proj e ct. intensive. seed. on se l ec t e d t o p i c s .. production. f rom. mo st. legume. Compared w i th c e r e a l s , grHin legumes are low y i eld ing. ( Hardy et a l . ,. 1977 ;. S inha, 1977) and t h e i r i mp o rt an c e i s declin i n g. re l a t ive t o c e r e a l s a s a s o u rce For examp l e , S i nha ( 1 9 7 7 ) s t at es. of. f oo d for human s and animals.. that over. t h e 2 0 years f rom. 1 9 5 2 to 1 9 7 2 , no i mp rov eme n t s to t h e av e r a g e y i e ld f rom l eg ume s su b s t an t i a l. o c c urred whe reas r i c e a n d whe a t. had. Howeve r , y i e l d i mpro ve ment s. occur. did. y i e l d increases.. in some l egume c r o p s. esp e c i a l l y so ybean s and p e a s.. Hardy. et. a l.,. ( 1 9 7 7 ) s u gge s ts t hat. t he. r a t e o f n e t p ho t o. syn t he s is i s a l imi t a t i on to yield in g r a i n l e gumes be ca u s e co2. en r i c hmen t of the c an o py re s u lt s i n s ub s t an t i a l y i e l d i n c r e a s e s in c ro p s su ch as s o y b e ans , p eas a n d pe an u t s .. T h i s hyp o t he s is. is suppo r t ed by Sh i b l e s et a l . , ( 1 9 7 4 ) a l tho u gh S p rent et al., . (19 7 7 ) , on f a irl y slen der evid ence , sugge s t e d t ha t a s s imi late was n o t a f a c t o r in Vicia faba yields.. Work i s the r e f o r e. u r g ent l y n e e ded. to b re ak thro ugh wh at. appears t o be a y ie l d barr i er in g r a in l e gume c r o p s .. It is. p l e as in g t o n o t e a c on s i d e r a bl e i n c r ease i n the amoun t of r e s e ar ch.

(25) 6.. bas i c physiology. the. work on. of. s o that our understanding of a lt h ou gh so. far,. no. legume s p e c i e s. many. prtlhlums. the. major solut itlllS. greatly. is. till�. to. i n rec e n t y e ar s in cre as e d ,. yiel d prob l em h av e. been found.. A.2. J)ACKGROUND. 1 NFOHMAT!ON ON. l.llPIN. of. T h e h i s t o r y and developmenl. r ev i ewed by v on Sengbusch (19 38),. this. in. 1970).. ,. is diverse were. There. The more import ant was the p r o v i d e d all. the species. s y s t e ms. a. .. l o n g es t. L.. per. i od. i s t h e olde s t. .. pr esently. attenlion. h is t o ry. and Afr i c a (56,000. (S i n h a. expan d e d. is. tram. for. t). the USSR. a w i d e range o f 197 6;. to. indigenous climatic. c. (W i ll i ams,. (SJO,OOO. t),. for the. mu tabilis. mo r e. a. 1 9 7 9).. today.. ( 1 59 ,000 t). Europe. ass u me d. to. be 1 97 2. 2,000 t on n es b u t. then would. a range df. ond i tio ns. L.. important. These are. production i n Australia since. Lup in s are. cultivated. been. pro d,,, · ed only. t h a t t i me, O c e an i a. agricultural. moden1. this ar e a bu t only r e ce n t ly. is noL. 19 7 7).. ,. w h ic h has. re gion. development i n t o. lupin. ,. evolution.. Suuth America.. this amoun t .. et a l .�. in. seed-producing crop plant.. on. At. useJ. The second cent rL'. lon g. lupin. Mediterranean-African. Produc t ion is c e n tr e d. f i gu r e s .. 1 97 7). be me n t i on e d only. two cl'nlres of. cu ltiva t e d sp e c i e s. Despite its. 1 97 4 ;. contains many species. and. Zbus is the sped es wll i t'11 llas. h as i t b e en r e ce iv i n g s op h i s t i c a t e d. (19 7 0;. Gladstones. review .. . The genus Lupinus. (Gl ad s to n e s. heen comp re h ensive l y. has. ( 1 9 7 6)) so theSL' aspects will. Hud s on e t al.� b rie f l y. lupins. have in c r eased. la t itu d es. (Gladston es,. to l e ra te s. and. 197 0;. Hudson. Sinha, 197 7 ), a l though adapt ab ility wi t h i n any one. s pe c i es is mo re l i m i t ed c o o l t empera te c l i ma t e s c l ima t e s (G lads t on e s, se ason o f a t l e a s t. .. Lupins and. are gro�1. as w i n t er. 1 9 7 0).. five mo n t h s. Best free. as summer. annuals. in s u b - t r op i c a l. lupin growing. from. a nn ua l s in. a reas have a. s e ri o u s mo is ture s t re s s.

(26) 7.. and du r ing tha t t ime , ha v e mean mon t h l y maximum t emp e r a tu r e s be tween 1 5 and 2 5 ° C ( G l a d s t one s ,. 1 970) .. S o i l s mu st b e f r e e. dra ining a n d can vary be tween m i l d l y a c i d t o s l i gh t l y a l k al in e wi t h some d i f f e ren c e s i n r eq u i r emen t s b e tween s p e c i e s ( G l ad s t on e s , 1 9 7 0 ; 1977) .. A. 2 . 1. LUPI N S EED QUALITY. S e ed q ua l i ty and f e e d va l u e f o r a r an ge o f an ima l s have b een widely r e p o r t e d ( e . g . G l ads tones , 1 9 7 0 ; W i thers e t a l . �. et a l . ,. 1978;. 1 9 75 ; Hud son ,. Hudson et a l . , 1 9 79 ;. 1972 ;. 19 7 7 ;. 1�76 , Hil l , 1 9 7 7 ;. H ove , 1 9 7 4 ; Hove. W i l l i ams 1 9 7 9 ) .. The mai n f e a t u re o f the seed is i t s h i gh p ro t e i n c on t e n t ranging be tween 2 8 % and 4 3% c r u d e p r o t e i n d epending o n Lhe s pe c i e s . Th e n i t roge n - f r e e e x t rac t le gumes ( 5 5 - 7 5%). (Hi l l ,. is. l ow ( 2 5 - 4 5 %) c ompar ed w i th mos t. 1977) .. The ma i n d i sadvan t age o f l u pin. seed i s a t h i ck hu l l wh i c h makes up 1 5- 2 5 % o f th e seed dry w e i gh t . Thi s h u l l c on tains much o f t he f ib r e b u t l i t t l e of the p ro t e i n s o t h a t i t s r emova l has the e f f e c t o f c ons i d erab l y improv i n g the feed va lue o f the s ee d .. A l k a l o i d is t he s ign if i can t t ox i c ma t e r i al. and on ce t h i s f ac t o r i s remov ed by t he use of swe e t c u l t ivars the seed. is. sa f e r. than. moa t. other. legume seeds (Hill, 1977;. Hud s on ,. 1 9 7 8 ) wi t h o u t the n e ed f o r f u r t h e r processing ( Hove et a l . , 1 9 7 8 ) .. Hudson ( 1 9 7 9 ) and W i l l i ams ( 1 9 7 9 ) b o t h be l ieve t h a t l up i n seed i s s u i t ab le f o r human c on s ump t i on and. it. b e s u i t a b l e f o r t h i s p u r p ose ( H e r z ,. Gross e t a l ., 1 9 7 6). 1 973 ;. has been s hown t o. b u t b e f o re i t can be wide l y used in human d i et f u r t h e r t ox i c o l o g i c a l studie s are p r obab l y r equ i re d ..

(27) 8.. A.2 .2. LUP I N S P EC I E S. De t a i l e d d e s c r i p t ion s o f. u. pub l ished by G l ad s t ones ( 1 9 7 4 ) .. extensive l y d e ve l o ped L . a ngus tij"o l ius L. ,. the species s t ud i ed f ore be given t o. some a t t e n t i o n. an d. t he m. in. in. is. <.JS. 1. s e l! cl and. • •. p r u g r a mme .. A u s t r a l i a i:l t t cl. being se l ected in S o u t h Ame r i c a a n rl a low alkaloid , n o n -s h a tt e r i n g. A. 2. 2. 1. Lttp i lltts. <l n g u s 1 i f < > I i. This s p e c i e s i s o ften. pod s. 1 949) .. ( Clads t o nes,. 1 9 7 0).. the. ttp. i n. c. s pe c i e s. ro p s .. been. These are. l u teus. t-lo s t. has. species have been. 3. on l y. r... w h i c h are also. a t t e n t i o n w i l l there. c:Ds e n t i rd i h a s received. L.. £, . rnu tabi l i s is. n u mb e r of. H tHl h i g h e r. r e f e r rL!d. Th i s. developed compa red with. l. c u rr e n t ly. o t h e r cen tres f or. s e ed yie l d .. tJS. a re. or b lu e l u p i n , a l t h ou g h t h e r e colour s ( A l l e n ,. L.. r e v i ew .. this. Wes te r n. nt. H ow e v e r. grown. L. ll l bu.;. i n th. n u mbe r. to a. b p e c i L! S o t he r. t h e n a r ro w - l e a fed lupin. as. w i de. range. o f p o s s i b l e f l ower. w a s r � l a t i v e l y l a te t o be. S P L' l' i e s. t md e r. c o nsideration. R e c en t l y t h i s s p e l' i es h a s b e e n i mproved by. selec tion f or red uc ed s ha t t e r i n g o f po d s , reduced ve r n a l i s a t i on more recent l y , d i se ase. re q u i r e me n t resis t a n c e. three cu l t ivars we r e r e l e a s e d. .. fi rs t t o have the comb i n a t i on. uf. wh. a l l ow i n g. i te flowers and seeds ,. e a r l i e r f lower ing , and. ( � l a d s t on e s 1 9 7 7 ).. U n i 1" h i t e l ow. ( r e l e a s e d 1 9 6 7 ) was t he. a l ka l o i d ,. seed and reduced sh a t t e r i n g ( C l ad s t on e s. I n i tial l y. 1 9 7 2 ).. ,. w h i t e flowers and. Uniharvest was. released in 1 97 1 and had increa s e d s hatt e r ing resi s t an ce but was o the rwise s i mi l a r to Uniwhit e .. l l n i c r op. t o Uniharvest except f or t he gen e ve rnal isa t i on req uirement. o f ten. Ku. ( re l e a s ed 1 97 3 ) is isogenic. wh i c h removes mos t o f t he. r e s 1 1 1 t in g. rela tive to Uniwhite and Un ih a r v e s t. i n ear l y fl owering. ( R ahma n and C l ads t ones ,. 1 97 2 ). I n a series o f 8 trials in We s t � r n Au s t r a l i a , Walt on ( 1 97 6 ) showed that Uniwhite was 4 - 6 2 % l mv e r i n y i e l d than Unicrop . Wit hers ( 1 97 3 ) and G a r s i d e ( 1 9 7 9 ). found. t h a t Uni c rop had only a. small advantage i n wetter and c o o l e r e nv i r o n me n t s .. Wal t on ( 1 97 6 ). at t ribu ted di f ferences be tween t h e c u l t ivars in his t rial s t o t he greater s usceptibi li ty o f Uniw h i t e t o s hat tering in hot . dry summers..

(28) 9.. The diff eren c e s b e t we e n l l n i h a rv e s t. Un i c ro p were sma l l excep t. and. in warm , s ho r t s e a so n d i s t r i c t s wh e n U n i c r o p had e a r l i e r f lowe r i n g (Gladstones ,. 1 9 72) .. Mo r e r e c e n t l y ,. two new c u ] t i v a r s h a v t� b e e n deve loped ,.,hi ch. hav e re s i s t a n c e t o g r ey l ea f s p o t ( .'> t c�lnphy l-i um ves1: ca r•iwnJ and o t he rw i s e. an t h r a c n o s e (C lome ::;e l la c1:ngu la ta ! b u t t o Uni h a r v e s t o r Un i c r op (. G l a d s t on e s ,. 1977;. L . angu s t ifo l i us i s t h e mo s t c o o l L o ] e r a n t. co n s i d e r e d i n t h i s t h e s i s ( G 1 a d s t o n e s ,. of. 1 970 ;. t h ey a r e s im i l ar. Anon ,. the t h r e e s p e c i e s. 1972. requ i r e s m i l d l y a c i d t o n e u t ra l s o i l s ( G l ad s t on e s ,. deep and f r i a b l e ( G l ad s t o nes ,. s ugge s t s a w id e r ra n ge o f. s oi l. 1 9 79 ) .. ;. 1977) .. 1 9 7 0 ) wh i ch a re. Howeve r , Co r b i n. 1977) .. It. t y p e s ma y be s u i t ab l e .. ( 1 9 7 8b ) Al though. e f f i c i en t a t u t i l i s i n g p h o s p h a t e r e s e r v e s c omp a r e d w i t h o t he r c ro p s ( G l ad s t on e s ,. 1 977 ). L.. ph o s p ha te and po t a s s i u m d e f i 1 970) .. Th i s spe c i e s h as b e e n. a. an';-JliS t ifo l i us c i en. · i e s t han L .. an d. n o n - s h atter ing p o d s. ( G l ad s t o n e s, 1 9 7 7 ) .. have been r e l e a s e d s i n c e 1 9 5 0 t h a t a g r i c u l t ure t h a n. p r e v i ou s. lut eus ( G l a d s t o ne s ,. s u b s i s t e n c e c r o p f o r over 3 , 000 y e a r s. and i t s e a r l y f o rm s h a d so f t s e e d s ( c f . L . angus tifo l i us ). i s mo r e s us ce p t i b l e t o. on e s .. are. Ul. A. n umb e r o f c u l t ivars. b e t t e r adap t e d t o mod e rn. t ra. ,. the main cu l t ivar u s ed. in t h i s s tu d y , was o r i g i n a l l y r el e a s e d f r o m a p r iv a t e German b re e d in g p r o g ra mme ( P f l u g s ) in. 1 9 5 0 b u t i t was s u b s e q uen t ly r e . s e l e cted i n We s t ern Aus t r a l i a hy G l a d s t o n e s ( G l a d s t on e s, 1 9 7 7 ) an d t\. i s b e i n g exten s i vely grown i n Eas t e r n Au s t r a l i a n S t a t e s ( J . S t rang p e rs . comm . ) .. L.. a lbus r e q u i re s h igher l ev e l s o f f e r t i l i ty and r a in f a l l. t h an. the oth e r l u p in sp e c i e s b u t h a s t h e h i ghe s t y i eld p o t ent ia l ( Gladstones,. 1 9 59 ;. 1 9 70 ) .. I t i s cha r a c t e r i s e d by a long period. f rom f l owe r in g to ma t u r i t y (G l ads t on e s ,. 1 9 5 9 ) wh i c h c an l imit i t s. u s e f u ln e s s i n areas w it h s h o r t p e r i ods o f w e a t he r s u i t a b l e f o r.

(29) 10 . ( l.u c a s ..J t . 1 ! . , 1 9 7 6 ) .. ma t u r i n g t h e c ro p by A l l en ( 1 9 4 9 ) not ripen. t o b e un s u i t ab l e. r e gu lar l y. charac t e r i s t i c .. ,. p r e s u ma b l y. g r ow s. Tt. po s s ib. t o l ow t e mp e r a t u r e s ( G la d s tone s ,. A. 2 . 2 . 3 L.. 1 970) .. l u t�us. has. been used. Med i t e r ran ean r e g i o n. as. ex ten sive l y a s a g a r d e n p l a n l A f ter about. 11 f. l. a11d. on. t i on s t han. i t s t o l e ran c e. gre en ma nu r e c ro p i n t h e. uf. i ts. c u l t i vu l ed. 1 970) .. of man y p a r t s o f Europe ( G l a d s t o n � s ,. level ( 4 2 % ). c on d i. d i f fe r i n g eco types. hun d red y e a rs. b e c a 1 1sc. h i gh l y. r e po r t s. �. I o ra g e. 1850 i t be ca me witl el y. A s a s e e d c ro p i t i s. wa rmer. s l i p; h t l y. l.ltJ> l n ). a. se v e r a. f or. t i ons as i t d i d. t o t h i s l a t e r ma t u r i n g. he�aus�. i l l l l ' I I S ( Y l ' l l tn.J. Lttp i ntJS. c on d i. Zea l and. N ew. v a r i ahl. ar�. ly. d ue. in. wel l. t h e o t h e r s pecie s h ut t he r e. Ft)r. a l bus was c on s i der ed. L.. d L· s i L.l b l e. a. ye l. n d was on ce u sed l ow s c e n t ed f l owe r .. on t h e ac i d soils. b c c a l i S e 'i t s s e e d prote i n. i s h i ghe s t o f t h e t h r � e l 1 1 pi n s p e ci e s co n s i dered here ,. with slightl y b e t t e r p ro t e i n Wither s e t a l . ,. 1 975) .. spe c ies i s u sua l ly. l. ( G l a d s ton e s ,. qua l i t v .. U n fo r t u na t e l y t h e. ow e r. t h an. fnr. L.. seed. yield. 1 970 ; of. this. . t ngus t ifn l i us an d L . a l bus. under Australian and New Ze a l a nd , · o n di t i o n s ( G l a d s t o n e s , Stoker,. 1 9 7 5 ) a l t h o u gh Pa l me r ( I Y 7 h ). bet ter for t h e no r t he r n N o r t h l s l a n d . L,. l u t eus t o l e ra t e s ac i d so i l s. i n d i c· a t e d. a11d. br ief. t h a t i t m a y be. p e r i o d s of w ater . logging bette r than the o t h e r t wo s p e c i e s ( G l a d s t one s , is also relat ively better on. poo r e r. 1 959 ;. 1972) .. It. s o i l s as i t i s s l i g ht l y more. toler an t o f low pho sphate and p o t a s s i 1 1 m l evels t h a n L . angus tifo lius ( an d p resumably L. a l bus ) 1 9 70 ;. ( G lad s t o n e s. Rahman and Gladstones ,. 1 974) .. et. ,·d . ,. 1 9 64 ;. G l ad s tones.

(30) 11.. THE DEV ELOPMENT O F. A. 3 .. A. 3 . 1. L EG UM E S E E D Y I E LD. P l an t S t r u c t u re a n d Deve l o p me n t. The d e ve l opmen t of s e e d y i e l d i n many i n te ract ing f a c to r s. Me ad l e y and. Mi l b o u r n. l n f l ue n c i t 1 g. g ra i n t l 1e. legumes i s complex wi t h. f i na l. n u t c · o Ptc ,. ,, ..:: e d. y i eld ,. ( 1 9 7 1 ) r � c ogn i s e d t wo st a ge s in t h e. development of seed y i e ld wh i ch. are : -. Vegetat i ve growth w h i ch p r ov i d e s t h e structure on wh i c h seed. (a). yie l d is u l t imately f o r me d .. The y. s ug g e s ted. po tent ial i ncreases wi t h v e g e t a t i v e growth rela t ionship w i t h n o d e n umbe r .. that repro d uc tive. l a rge l y. Flowering and podf i l l ( r e p rod u c t i v e g rowth) .. (b). because o f i t s. T rans fer o f. ass imi late f i xed i n the p r ev i o u s p e r i od i s l i m i t ed s o yie l d i s dete rmi ned l arge ly by d r y we i gh t a c c u mu l a t e d i n t h i s s tage .. workers (e . g . Herr i dge and P a t e ,. 1977;. d i vide t he second s tage f ur th e r i n t o. f. growth plus po d f i l l and ma t u r i t y .. An optimum balance of v e get a t i v e. important to ach ieve h i gh. se ed. 1 9 7 7 ) sub. l o we r i n g and ear l y fru i t. to. r e p r od uc t iv e g rowt h is. y i e l d ( Ad ams , 1 9 7 5 ;. 1 9 7 7 ) b ut the spec i f i c b a l an c e h a s no t. There i s some ev i dence i n Vi cia. Sun�e r f ield,. Man y. b e en. Su�er f ie l d,. determ ined (S inha ,. 1977).. f'aba f or exam p le t hat the amoun t. o f vegeta tive growth i n no rma l p l an t s i s exce s s ive (Chapma n and Pea t ,. 1 9 7 8) .. Adams ( 1 9 7 5) described. t h r e e t y p e s o f p l an t developmen t :. (a) where vege ta t i ve and r e p r o d u c t i v e phases d o not over lap i. e . vegetative deve lopment cea ses a t the o n s e t o f f l ower i ng . are res tri c ted i n number but not s i z e .. Leaves. Examp le o f this type o f. g rowth c a n b e found i n some ga r d e n pea cu l t i vars developed for process ing and some ear ly soy b e an cu l tiva r s .. Th is is the type o f. growth genera l ly re ferred t o as determ i na te (Eg l i and Legge t t ,. 1 9 7 3) .. Sinha ( 1 9 7 7 ) sugges ts tha t the sh o rt vegetat ive growth o f these c rops reduces their yie ld po te n t i a 1 .. Their small p l an t s i ze however,.

(31) 12.. usua l ly a l lows the smalle r y i e l d pe r p l a n !. p l ant i n g dens i ty .. I n a r e a s o f unce r t ai n rai n fa l l ,. p l a n t s may be less r e l i ab l e ( S i nha , (b). t o be o f f s et b y gre a t e r. ] q 7 7) .. Wh e re v e g eta t i ve g r ow t h c o n t i n u e s f u r· a bout. the ons e t. of. Mat1 1 ri t y o f. f lowe ring .. a n d the y a r e i n de t e rminat e , and vining pea cult i. va rs .. o f 4 4 days fo r. 2 0 - 30 d ays. Exam p h, s gi v e n are many P . L u p i n s t l s ual l v f i t. F o r e xampl e,. after. L h e se p l a n t s i s u s ua l l y l a t e. the p otentia l t o c o n t i nue v e g e t a t i v e g row t h a f ter f l owe r i ng .. d e t erminate. A t k i ns. d. t his c a t eg ory bu t have. the 20-30 days. be y o n d. o. 7 . ,. v u l garis. '. ( 1 9 7 8 ) no t e d a p e r i o d. f l owe d ng and e a r l y r o d f -IJ l d u r i ng wh i ch t ime. v e g e ta t ive p ar t s grew v i g o r o u s l y .. (c). Wh e r e t he vegeta t i ve phas e i s J o r t g , n e a r the end o f wh i c h a. l ong perio d o f f l ow e ring and p od s � L. c o mme n c e s. b ut l im i te d ove r lap o f. v e ge tative. an d rep r o duc t i ve g r ow t h L J C L" t t r s .. s oyb e ans .. S om e l ur i n cul t i vars h a v e t l 1 e charac t er i s L ic o f a l on g. vege t a t ive p e riod ( e . g .. L.. a l buH ). but. a re. e .R.. N o r th Ame r i c an. m o r e i nd e t e rminate than. imp l ied i n this de sc r i ptio n .. A f eature o f t yp e ( b ) and,. to. a s ma l l e r. e x t e nt ,. ( c ) p l ants. is. the p o t e nt i a l that e xis t s for compe l i t l o n betwe en vegetat i ve and r e p roduc t i ve grow t h ( E g l i and L e g g t.!l t , Gre enwood e t a l . >. 1 9 75 ;. i n r e ducin g p o d s et .. Far d n g t u n ,. 1973;. S hi b l e s. et a l . >. 1 9 74 ;. 1 9 7 6 ) wh i ch ma y b e a f a c t o r. The l o ng veg e t a t l ve p e r i o d o f t ype ( c ) may b e. a d i sa d va n t age as i t r e d u c e s t h e chan c e o f t he s e ns i t i v e f l owe r i ng and pod s e t o c c u r r i n g und e r favour a b l e c ondi t i o ns ( Adams , 1 9 7 5 ;. S inha ,. 1 9 7 7 ) wh i ch l imits the i r a b i l i ty t o r e a l i s e the h i gh p o t e n t i a l f at·. s e e d y i e l d wh i c h exists i n th e i. r. can o p y s t r u ctur e. ( Adams ,. 1975) .. Pe r ry and Poo l e ( 1 9 7 5 ) sta t e tha t the i nd e t e rm i na t e g r ow t h h a b i t i n l u p i n c a n f u l ly exp l o i t the e nvi r onmen t b e c ause i t c a n va ry i t s g r owth p e r i od and that s e ed y i eld. i s n o t so d e p end ent on ear ly. c ont r a s t s w i th. g r owt h ;. a s i tua t i on wh i c h. howeve r ,. sugge s t s t hat the abi l i t y o f. c ereal s .. l up i n s. Ga r s i d e ( 1 9 7 9 ). t o u t i l i s e t h e ava i lab l e. season i s a c a u s e o f the large vari ation i n y i e l d ..

(32) 13.. Third o r d e r l ate ral b ra n c h. �. Second order later a l b ranch. F i r s t ord e r lateral branch. branc h n umber .-+ 1 : 3. M a i n stem. Fig. A . 3 . 1. S chema t ic l ay o u t o f a nd number i ng. a. l u p i n plan L. sh owing. o f t h e b r a nc h orde r s .. t h e pos i t ion.

(33) 14 .. De t a i J e d d e s c r i p t i o n s o f t l 1 e. p l an t w i t h empha s i s o n L . aruJUB t ij"o l iwJ. a n d Gre enwood ( 1 9 7 5 ) ,. are. G r e enwo o d e t a l .. ( 1 9 7 5 ) , Fa r r i n g t on ( 1 9 7 6 ) a n d. et �l.. Re e v e s. the ma in s t e m. an d f u r t h e r. n um b e r. A v a ry i n g. each. of wh i c h c a n. Fa r r i n g t o n a n d. I n b r ie f ,. ( 1977) .. w i t h a t e rmin a l. s tem). l a t e r a l b r an c h e s a r i se from. p rnd u c e. fn. l at e r a l b ra n c he s .. o r d e rs are deve l op e d a s. of. p r o v i d e d b y F a r r i ng t on. ( 1 9 7 5 ) , P e r r y and P oo l e. i n it i a l g rowt h i s by a s i n g l e s t em ( main in f l o re s c e n c e .. o f t he ann u a l l u p i n. s t ruc t ure. a. t e rm i n a l i n f l o r e s c e n c e. t h i s wa y a s e r i e s o f b r an ch in. i 1 l tt s t ra t e d. A. 3. l.. Fi g .. ( 1 9 7 5 ) a n d P e rr y a n d P oo l e ( 1 9 7 5 ). G r e e nwo od. u s e d i f f e rent s y s t e ms f o r d e s c r i b i n g t h e v ar i o u s b r a n c h o r d e r s . The s y st e m proposed b y F a r r i n g t o n a n d G r e e n wo od. ( 1 9 7 5 ) i s the. more. c o m p r e he n s i ve but is more c omp l e x t h a n r e q u i r ed in the work rep o r t e d here s o a mod i f i c at i o n o f. and Poo le ( 1 9 7 5 ) has been A. 3. 2. C o mpon e nt s o f Le g u me Seed y i e l d. in. a do p t e d. is bui lt. up. t u rn a r e d et e r m i n ed by. f ac t o r s .. t h e nome n c l a t u re us ed b y P e r r y. ( see. Seed. Y ield. by. se r i e s. a. A. 3. l ) .. Fig .. uf. yield. componen t s wh i ch. a combi nat i o n o f p l ant a n d e nv i ronmen t a l. A d ams ( 1 9 7 5 ) d e s c r i b e d t h e r e l a t i on sh i p o f t h e s e. c omponent s t o p l ant st r u ct u r e ( F i g . A . 3 . 2 ) .. The d ev e l o pmen t o f. l u p i n y i e l d compon e n t s h a v e recen t l y b e e n d i s c u s s e d b y s om e. Au s t r a l i an wo r k e r s w i th e mpha s i s on P e rr y ,. 1975 ;. Pe r r y a n d P oo l e , +-. Po d s / P l an t. L. l975 ;. anr1us t i.fo l-i us ( e . g .. G a r s ide ,. 1 979) .. Tot a l N o . No . l e a ve s +... No . b r a n c h e s no de s / p l an t p e r p l a nt. t. Y i e ld. No . l o n g i n t e r no d e s. S e ed s / P o d. ". t. Pod �. /. l e n g th. P l a n t He i gh t. /. t. Leaf s i ze. We i gh t / S e e d. Fig A . 3 . 2. In t e rnode. \. D i a g rama t i c r e p re s e nt at ion o f d i re c t p a t h s i n f l u e n c e o f s e ve r a l p l an t u p o n y i e l d. s t ruc t u r a l. ( Ad ams ,. 1 975) .. comp on e n t s o f. of the.

(34) 15.. The n u mb e r o f p o d s. pe r. un i t a r ea o r p e r p l an t i s u su a l ly the. ma in c omp on e n t t o in f l u e n c e y i e l d I shag ,. 1 9 7 3a , b. and Hunge r ,. ( Wa l l a c e. The. Adams , 1 9 7 5 ) .. 1966 ;. n umb e r o f pods r e qu i re d f o r. h i gh y i e l d i s dep en d a n t on a sa t i s f a c t o r y p l an t s t ru c t ur e whi ch i n f l uen c e s t h e n umb e r o f f r u i t i n g 1 975 ) .. In l u p ins t h e pod n umb e r. n od e s .. is. ( l shag ,. d e t e rmi n e d. 1 9 7 3a ;. Adams ,. l a rge l y b y t he. numbe r o f l a t e r a l b r anches wh i c h i n f l ue n c e s t he numb e r o f i n f l o r e s cences f o rme d .. ( W i the r s ,. 1973 ;. Gars i d e ,. uni t s h ave b e en d e t e rmined b y. the. s t r u c t u re , t h e r a t e o f. p l an t. ab s c i s s i on o f f l owe r and. s ma l l. pod number ( G abe lman and. W i ll i am s ,. O n c e the n ur:b e r o f f lo r a l. 1 9 79 ) .. pods. a n i mpor t an t d e t e rminan t o f. is. 1 96 0 ) .. Th i s. a s p e c t wi l l be. d i s c u s s e d f u r t h e r i n S e c t i on A . 3 . 3 .. P o d n umbe r i s usu a l l y t he. f i rs t. seed. y1. e. l d compon e n t t o b e. in f l ue n c e d b y t h e env i r onmen t a n d c u l t u r a l pra c t i ce s ( Benne t t e t a l . , 1 9 7 7 ) and i s very se n s i t i v e t o. st. re ss i n. th i s ready adj u s t men t , o t h e r compon e n t s p e r p o d and seed we i gh t , Se i t z e r an d Evan s , 1 9 7 3 ) .. t end to Seeds. be. t he. p lant .. Be cause o f. y i e l d , v i z numb er o f seeds. of. l � s s vari ab l e ( I sh a g ,. pe r. 1973a;. pod and seed w e i gh t are ,. howeve r , c apab le o f c on s i d e r ab l e c om p e n s a t i on de pen d i n g on the numb e r o f pod s s e t on the p l an t ( Adams ,. 1 96 7 ) .. I n c r e a s e s in the. number o f see d s per pod and / o r s e e d w� L g h t can o c c ur in r e s p on s e t o l ow p o d numbe rs o n a p l an t ( Ch u n g 1 9 7 3a ;. Ndun guru et a l . ,. 1 9 78) .. G r e e nw o o d e t a l .. in L , angus tifo l i us t ha t a b o u t 1 / 6 o f. ma t ure s e ed ind i c a t ing p o t e n t i a l. Co u l den , 1 9 7 1 ;. and. t he. I s h ag ,. ( 1 9 7 5 ) noted. ov u l e s d i d n o t produce. for f l e x i b i l i t y in the n umber o f. seeds p e r pod .. S e e d s i z e i s d e t e rmin ed by. gen e t i c. and. env i ro nme n t a l inf luence s. and i s usu a l ly nega t iv e l y c orre l a t e d w i t h s e e d numb e r ( Shib l e s et a l . , 1974) ,. Pro t e in c on t en t o f the seed d o e s n o t v a ry grea t l y ( S e i t z e r. and Evans , 1 9 7 3 ;. Woodwar d and B e g g ,. 1 9 7 6 ) a l t hough Far r ington. ( 1 9 7 6 ) n o t ed l ower ni t ro gen c o n ce n t r a t i on i n t h e l a t e s t d eve loped. s e e d s o f l u p i n s . Th e s e we re , h oweve r , l i ghter i n we i gh t and n o t f ul ly d eve loped ..

(35) 16.. l up in s. In c o mp ared. y i e l d c o m p o n e n ts on t h e mi.l i n s te m ten d t o b e s t ab l e b ra n c h e s .. Seed. n umb. · ·. r. p e r p od and. we i gh t u s u a l l y r e d u c e a s l a t e r a l b ran ch or d e rs i n c r e a s e. seed. ( G re enwo o d e t a l . , and. l a te r a l. w i th tho s e on. Hi l l ,. 1 9 7 8b ) .. 1 9 75 ;. Pe r r y ,. 'l11 e e f f e c t. on. the l en g t h o f t i me be tween f e r t. seed. i1i. d e v e l o pm e n t o f. compon e n t s a r e " t o. some. e x t en t. He r b e r t. ra p i d ::; e ed f i l l wh i ch. and. la t e r a l b r a n c h e s ( P e r r y , 1 9 7 5 ) .. c o m p on e n t s. yie ld. 1976 ;. w e i gh t m a y b e a f a c t o r o f. s a U on. te n d s t o be sh o r t e r f o r h i gh e r o r d t.� r. Th e. F u r r i n g ton ,. 1 975 ;. i s s e q uen t i a l a n d t h e. i n t e rd e p e n d e n t. i n th e i r d eve l o pmen t ". (Ad ams ,. 1 96 7 ) .. me a n s. s i z e c a n take a l te rn a t i v e p a t hw a y s d e p e n d i n g o n e n v i ronmen t a l. co n d i t i o n s .. i n t h e d e v e l opmen t o f y i e l d c omp on en t s. "P l as t i c i t y ". a. Th i s he l p s t o ma i n t a i n. s t ab l e y i e l d l eve l .. The. ten den c y f o r compon en t s to c ompen s a t e and to b e n e g a t iv e l y c o r r e l a t e d ,. p l u s the t en d en c y f o r p o t e n t i a l s i te s n o t t o be r e a l i s e d , ind i c a t e s. a li m i t a t i o n i n a s s i m i l a t e ( A d a m s ,. Dunphy a n d H a n w a y ,. 1 9 76 ;. S h ib le s e t a l . ,. 1 96 7 ;. H a r d y e t ,d . , 1 9 7 7 ). .. 1974 ;. S t i mu l a t in g ph o t o. s y n th e s i s c a n i n c re a s e y i e l d c o m p o n e n t s ( H a r dma n an d B r u n ,. Schou e t a l . ,. A.3 . 3. 1971 ;. 1 9 78) .. F l owe r i nJL a n d P o d s e t. T i mi n g o f s p e c i e s and. c. f l or a l. in i t i a t i o n. u l t i va r .. P l an t s. can. f ac t o r s - u s u a l l y v e r n a l i s a ti o n , t em p e r a t u r e s .. Th i. Kr i e d emann ( 1 9 7 6 ) .. can. be. be. c omp l e x d ep en d in g on. v e ry. r e s p on s i ve to o n e o r more. and / o r i n c re a s i n g. p h o t o pe r i o d. s t op i c h a s b e e n. r e v i ewed. Lup i n s u s u a l l y h av e a. by Le o p o l d a n d. h i gh. verna l i s a t io n. req u i r emen t an d a we a k p ho t o p e r i o d re s po n se ( S e e S e c t i on A . 5 . 3 ) .. B a s e d on the n umbe r o f po t en t ia l f o r h i gh. yields. f l o we r s. p roduced ,. ( G r e e mv o o d. et al . ,. l e gumes have a. 1975;. S in h a ,. 1977). b u t u s ua l l y on l y a sma l l p r o p o r t i o n o f f l owe r s p r o d u c e d s e t ma t u r e p o d s a l tho ugh a l l f l ow e r s a p p a r e n t l y have t h e ab i l i t y t o s e t pods ( van S t even in c k , 1 9 5 7 ) . F r an c i s et a l . pods ,. For a. r ange. of. s. i te s i n We s t e r n Au s t r al i a ,. ( 1 9 7 1 ) f o u n d tha t o n l y 6 - 3 3% o f l up i n f lowe r s s e t. McAl i s ter and Ko b e r ( 1 9 5 8 ) a n d Adj e i -Twum a n d S p l i t t s toe s s e r.

(36) ( 1976) ad j. rega r d. the. loss. of. a n d p od s a s a mec han i sm for. f l ow e r s. u s t i n g t h e r e p r o d u c t i ve l o ad. to. t h e capac i t y of. t he p l an t t o. s u s ta in i t . Th e r e. wh i c h. to. be. !TlCl. a l one. or. in. appears. can. ac t. c a uses. ny. o f f l ow e r a nd p o d a bs c i s s i o n. c om b i n a t i on. S inha ( i 9 7 7 ) l is t s the. .. f o l l owing :. 1.. L i m i t e d p h o t o s y n t ha t e. 2.. L i mi t e d n i t r o g e n ava i l a b i l i t y. J.. Re d u c e d l i gh t. 4.. C a n o p y t em p e r a t u r e. 5.. H o rmo n a l. 6.. P o o r g a s e x c hange. 7.. H um i d i. 8.. S o i l and wa t e r f a c t o r s. Many o f a s s i mi l a t e. i n t C' n s i t y i n t h e c a n o p y. t y i n t he c a nopy. thes e. f a c t or s a r e i n t e r - r C' l a t ed a n d c a n i n f l u e n c e. ava i l a b i l i t y . t o be. n i t r o g e n seems. i n t he ca nopy. t he. I n t e r na l s u p p l y o f c a r b o h y d ra t e a n d / or mo s t w i d e l y f a v o u r e d ba s i c c a u s e o f. a b s c i s s ion ( Ad d i c o t t and L y n c h ,. C oo p e r e t a l . J. 1978;. S t ewa r t e t a l . J. mov e m e n t. Eg l i a n d Legg e t , 1 978) .. 1 974;. can. at. et a l . J. 1 97 5 ;. N d g u n gu r u e t a l , J. t em p e r a t u r e , a i r. f l owe r i ng a n d e a r l y p o d d e v e l opme n t. s u b s t a n t i a l l y i n c r ea s e p o d n umbe r s ( H a r d ma n and. S chou. I n t r a -p l a n t. L 97 6 ;. I n c r e a s i ng l i gh t ,. o r c o 2 c on c e n t r a t i o n. o f s oy beans. Brun ,. 1 976 ;. G r e e n w oo d. L 9 5 ') ;. et a l . J. c o mp e t. 1 978) .. i t i o n f o r a s s i m i l a t e be tween t h e i n f l o rescence. an d r a p id l y g r owing l a t e r a l b r a n c h e s h a s o f t e n b e e n ment i oned as the cause o f absc i s s i on i n l u p i n P e r ry and P o o l e ,. ( 1 957 ,. 1 9 58 ). 1975;. ( G r e e n wood e t al . J. H e r be r t ,. He d i f f e r e n t i a t e d. of. b e t we e n. c a u s e d b y la t e ra l s t em g ro w t h ). area ) ,. P e r ry ,. 1975;. Ea r ly work b y van S t ev e n i n c k. i n d i ca t e t h a t a h o rm one may be i nv o lv e d i n ea r ly. a b s c i s s i on b u t t hat t h e s u p p l y l a t er ,. 1 9 7 7c ) ,. 1975;. and. as s. i m i l a t e may be m o r e imp ortant. f l ower a b s c i s s i o n ( a p p a r en t l y p od. absc i s s i on ( r e l a t e d t o l e a f. y.

(37) 18.. Ab s c i s i c a c i d has b e e n imp l i ca ted a s a n i mp o r t an t c au s e o f ab s c i s s i on in w a t e r - s t r e s sed. L.. (Porte r ,. l uteus. 1977) .. It is. f ound i n i nc r ea s i n g q uan t i t i e s i n a b s c i s i ng p o d s o f t h i s s p e c i e s. a n d i n pod s , s e e d and l e a v e s o f L . a lbus p la n t s s ubj e c ted t o. w a t e r s t re s s ( H oad ,. 1 978) .. Z u c c o n i ( 1 9 7 5 ) and P o r t e r ( 1 9 7 7 ). b e l ieve t ha t t he p re s en c e o f a b s c i si c a c i d i s a n ef f e c t o f s t r e s s r a t h e r than. a. c a u s e o f ab s c i s s i on .. S ubhad r ab an dhu e t a l .. ( 1978). d id n o t f i n d a c on s i s t e n t rel a t i o n s h i p be tween ab sc i s i c a c i d and abs c i s s ion i n P . vu lgaris .. E t h y l ene has a l s o b e en i mp l i ca t ed i n. a b s c i s s ion ( McMi cheal e t a l . � 1 9 7 3 ) .. Ad d i c o t t a n d Lyn c h ( 1 9 5 5 ) s ug ge s t s t ha t i n su f f i c i en t c arbo hydr a t e can c a u s e emb r y o abo r t i on whi ch r e d u c e s aux i n s u p p l y t o the ab s c i ss i on zone thus causing ab s c i s s i on .. Wa t e r s t re s s c o u l d. cause ab s c i s s i on by r educ i n g carboh y d ra t e s up p l y and / or b y d ire c t l y p r even t ing emb ryo g r ow t h , t h e r e b y c a u s ing emb ryo d e at h ; o r b y s l owi n g auxin p r oduc t i on .. S e e d s a r e a c t ive p roduc e r s o f. h o rmon e s wh i c h i n f l ue n c e s a s s i m i l a t e s u p p l y ( Harvey , 1 9 7 7 ) s o t h a t c e s s a t i on o f growth c aused by wa t e r s t r e s s may a l so reduce ho rmon e p r od u c t i on w h i c h i n tu rn , may r e d u c e the carbohy d r a t e supp ly to t h e p od , r e s u l t i n g in emb ry o a b or t i on .. on the s t ag e o f growth o f each f l owe r / pod . ( 1). Ra p id l y d ev e l o p i n g young p o d s. (2 ). Un o pened f lowe r s. (3). Opened f lowers. (4). Youn g f e r t i l i s e d ovu l e s in d eve l o p in g pods. ( 5). Freshly p o l l i n a t ed emb ry o s and very young p od s .. When compe t i t i on f o r ass imi l a t e s b e g i n s ( i f tha t i s t he c a us e ) , ( 5) i s ab s c i s e d f i r s t , f ol lowed by ( 4 ) e t c , depen d i n g o n the dura t ion a n d ex t e n t o f the shor t age .. Adam s also sugge s t e d that. each n u t r i t i onal un i t t en d e d t o b e s u p p l ied equa l l y �-1i t h a s s imila t e . Howeve r , b e cause e a c h un i t was a t a d i f f e r en t s t a ge o f d ev e lopmen t , each w o u l d b e a f f e c t e d d i f f e ren t ly ..

(38) 19.. Max im i s ing s e ed y i e l d may t he o r e t i c a l l y b e ach ieved b y minimi s i ng ab s c i s s i on o f f l owe r s a n d 1 960) .. pods. ( Gabe lman and W i l l i ams ,. Thi s c on c e p t , howev e r , ove r lo o k s t he po s s ib i l i ty t h a t t h e. p l an t ma y a lr e ad y b e o p e r a t i n g w i t h i n envi ronme n t a l con s t ra i n t s by. and increased p o d s e t may b e o f f s e t y i e ld c ompon en t s .. l ower p ro d uc t iv i t y o f l a t er. I t seems un l i k e l y t ha t y i e l d improvemen t wi l l b e. a c h i eved b y work ing o n imp r ov ing ab s c i s s i on a l one i f the r a t e o f. a b s c i s s ion i s a r e su l t o f t h e in t e rn a l and ex t e rn a l env i r onmen t o f t h e p l ant .. A. 3,4. Pod and Seed Deve l opme n t. S inha ( 1 9 7 7 ) l i s t s 2 t y pes. of. ( a ) where t h e f r u i t wal l comme n c e s. f r ll l t. d eve l o pmen t : -. rap i d. grow t h f i r s t f o l l owe d b y. s e e d - t y p i f i e d by p e a s o r ch i c k p e a s .. Th i s t yp e o f d eve l opment. a l s o o c c urs in l up i n ( G ree nwood e t a l . � 1 9 7 5 ;. Farr ing t on ,. 1 976 ;. H o c k i n g and Pa t e , l 9 7 7 ) . ( b ) where p o d and s e ed deve l o p t n g e t h e r .. Examp l e s a r e mung b e an s. and cowp e a s . The grow t h o f t he pod and s e e d h a s. been. d e s c r ibed in d e t a i l f o r a. numb e r o f le gume s pe c i e s ( e . g . F l i nn a n d P a t e ( 1 9 6 8 ) f o r P . ar-vens e ; Car r an d S ken e ( 1 9 6 1 ) and O l i k e r. -vulgari s ) .. et. al.. ( 1 9 7 8 ) f o r Phaseoulus. Genera l l y , s e ed gr owt h i s i n i t i a l l y s low c o i nc i d ing. with rap i d p od. gr ow t h and t h en g r ows r ap i d l y .. O f t en a shor t " l a g. phas e " o c c ur s wi t h a sho r t p e r i o d ( 3 -4 d ay s ) o f s low g rowt h f o l lowe d. by a f u r t h e r p e r i o d o f rap id g r ow t h .. Dur ing th e l a s t per i o d of. rapid g r owth the pod u sua l l y lo ses dry we i gh t and n i t rogen an d mos t o f the pro t ein r e serves in t h e s e e d are a c c umu l a t ed .. Hocking and P a t e ( 1 9 7 7 ) comp a r e d t h e deve l o pmen t of P . sati-vwn L. angustifolius and L. albus .. Th e r e were s im i l a r general p a t t e rn s. o f pod , embryo and t e s t a d ry we i gh t a c c umu l a t i on a l though r e l a tive t iming of the r a p i d ac cumul a t i on r a t e v aried .. L.. albus had a. con s i d e r ab le lag per iod be f o r e t h e pod or emb ryo commenc e d linear growth c ompared wi th the o t her sp e c i e s .. P,. sati-vum commenc ed p o d. growt h a n d dec l ined i n d r y mat t e r e a r l i e r t h an L . angustifo lius.

(39) 20 .. a l t hough. e mb r y o. was l o s t b y. g row th. pod and. tes ta. r a p i d growth p h a s e in. bu t. e s t ima t e d t h a t p o d a n d t e s t a. 2 6 % f o r L. a l bus .. P.. F l inn. ea r l i e r. we i gh t and 2 3 % o f n i t r o g e n. t h i rd. i n P.. t i me .. Dry ma t te r. o f t h e emb ryo. sativwn .. l 7% f o r L.. up. I t wa s t o 1 3 . 5%. angus t �' fo li us an d. (1 9 6 8 ) e s t ima t e d 1 9 % o f s e e d d r y. Pa t e. r e q u i r emen t. b een made a v a i l a b l e f r om. pod ,. te s t a. The l i n e a r pha se o f. s e ed. i ncrease. r a t e wi t h i n a s p e c i e s. s ame. hav e c on t r i b u t ed. cou l d. sa t i v wn , and. t he. l ast. t he. d u r i ng. l up i n. o f t h e s e ed we i gh t f o r. abo u t. c omme n c e d. o f the embryo c o u l d have en. and. has. d o s p e rm o f P . ar vens e .. a. r e la t iv e l y c on s t an t. l 9 7 lb ;. ( H a nw a y a n d W e b e r ,. S i nc l a i r and d e W i t ,. 1 9 7 6 ) but d i f f e r e n c e s in the t i mi n g o r t h e commen ceme n t o f t h i s. phase c a n vary b e t w e e n. c u l t i va r s. v a r i a t i on c an o c c u r i n the r a t e and Ko l l e r. ,. and Hanwa y ,. 1 976 ;. S i n c J a i r and de l�i t ,. b e t ween c u l t i va r s o r. env i ro nmen t s .. f o rmed p o d s o f soybean. noted de. ma y b e. lup i n s . S e e d s o f t h e ma in s t e m o rd e r l a t e r a l b r a n che s. A.4. W eber ,. 1 9 7 lb ) ;. n on - l inear phase ( Ka p l an. 1975;. Eg l i et a l . �. 1 9 76 ;. 1978). d i f f e r e n c e s i n s ee d s i z e and /o r y i e ld. to. Egli and L e gg e t ( 1 9 7 3 ) have. al . �. t he. of. and. 1 9 7 4 ) o r i n t he d u r a t i o n o f t h e l inear pha s e ( Dunphy. a l l o f wh i c h c a n g i ve d s e. et. ( H a n wa y. H a mv a y t ha t. l ayed. .. the. and. Web e r ( 1 9 7 l a ) and. d ev e lo pme n t of e a r ly. S i mi l ar. i n f l o r e s c en c e. pat terns o ccur in. an d t h o s e o f t h e f ir s t. tend. to. d e v e lop t oge t h e r ( Gr e enwood. ECONOMY. OF. G RAIN LEGUMES. h o we v e r ,. P e r ry , 1 9 7 5 ) .. THE CARBON AND N I TROG E N. C a rb on and n i t rogen p ro v i d e t he b a s i c ' bu i l d ing b lo ck s ' o f y i e l d s o a n un d e r s tand i n g o f t he i r u t i l i s a t i on i n t h e p la n t i s i mp o r t an t f or the i n t e r p re t a t i o n a n d e x t er na l fa c t or s .. of. t h e p l ant ' s re a c t i o n t o i n t e rn a l. I n a t t e mpt i n g to ma xim i s e s ee d p r o du c t ion. we are t ry in g t o man i p u l a t e t he c a rb o n and n i t rogen e conomy t o a ch i ev e t h e max i mum amount o f s e ed p o s s i b le w i t h i n t h e env i ronment a l cons t r a i n t s . (a). Two a s p e c t s need t o b e c o n s i d e r e d : -. t o t a l a s s im i l a t i on o f b o t h n u t r i e n t s a s th i s wi l l d e t ermine. t h e m ax i mum p o t en t i a l o f seed p ro d u c t i o n .. Th i s is l a rge l y a func t ion. o f e nv ir onme n t and manageme n t wh i c h w i l l be c o ns i d e r e d m a i n ly i n S e c t ion A . 5 ..

(40) 21 . (b). d i s t r i b u t i on a n d. of. t h e max imum p r opor t i on i n c or p ora t e d. into. t he. seed. t he. Ea ch nu t r i e n t. o f b o t h n u t r i e n t s t o ens ure. r e d i s t r i b u t i on. wi l l. nut. t o t rJ 1. r i e n t s a s s im i l a t ed b e c ome. .. i n j t i a J ] y be c ons i d e red s e p a r a t e l y a s. p a t hway s a r e d i s t i n c t l y. cl i f f p r e n t. their. a r e c lo s e ly l i nk e d. a l t h ough t hey. a nd i n t e rd e pe nd e n t . A. 4. 1. Th e. E c o n omy. Carbon. f un c t i on. The i mp o r t a n t. t lte. p r i or t o f l ow e r i n g i s f l owe r i n g a n d. s e ed. c a r b on. 1959). to the me r i s t e m. and. most. of. l ea v e s a r e e xpor t in g a s s im i l a t es. wh i c h. 1 911 5 ;. Ove ra l l , pa t t e rn s o f a c t iv i t y a n d ( s in k s ). p os i t i on. in r e l a t i o n. (Ne l s on ,. of. to. on t h e d i s t ance a n d. ra t e ,. the. va. pos i. t hese. d i re c t i o n. As s i m i l a t e. 1 96 6 ) .. s t or ed i n the s t em ( C a r r. in. and. However , a s the l e a f the. movemen t o f e xp or t e d. 1 de f i ne d .. l owe r. 1 96 6 ;. Usua l ly ,. l owe r. s t em a n d r oo t s y s t e m. Wa r d l a w ,. 1 96 8 ) .. a r e l a rge l y d e t e rm i n e d by t h e. r i ou s orga n s r eq u i r i n g a s s im i l a t e s t i on. o f t h e supp l y o r g an ( s ourc e ). p a t t e rns h owever , of. t r a n s l o c a t i on. va s c u la r l i n k s be tween pa r t i c u l a r Porter ,. c a nopy ,. t he. m o v e me n t. the. Wi thi n. 1 963) .. n i t i a l l y i s t ra n s p or t e d upwa rd. 1 ess we1. l eav es are c omm i t t e d t o s u p p l y i n g Cor y ,. i. t he. in. a s s im i l a t e be c om e s c omp ·1 e x a n d ( B i d d u lph a n d. a f ra mewo rk on wh i ch s a t i s f ac t o ry. n ew l y d e v e l o p i n g l ea ves .. ma t ur e s and i t b e c om e s l owe r. ( f or s e ed p r oduc t io n ). t r1 k e p l a c e .. can. expa n s i on ,. (Tha ine et a l . �. ass imilat ion. h u i ] rl i n g o f. d e v e l opm e n t. ful l. Even be f or e. of. s on rc es. e xc e s s Pat e ,. t h e r e a r e r e s t r i c t i on s a s d e t e rm i ne d b y t h e. a n d s i n k s ( N e l s on ,. o f r e q u i reme n t s c a n be. 1 9 6 7 , Ward l aw a n d P o r t er ,. 196 3 ;. 196 7 ) .. In many g r a i n l e gume s , mos t o f t h e c a rbon i s f i xed by t h e p l a n t a f t er t h e c ommenc eme n t o f f l owe r i ng ( Ea g l e sham e t a l . ,. 1 9 7 8 ) and t h i s. i s t h e ma i n sou r c e o f c a r bon f o r s e e d y i e ld (Me a d l ey a n d Mi l b o u rn , At k i n s et a l .. o f L . a lbus wa s. ( 1 9 7 8 ) n o t ed t ha t o n l y 6 % o f t h e t o t a l n e t p h o t osyn t h e s i s f i xed. be f o r e f l oweri ng .. used i n the l ea f l e t s ( 2 7 - 3 0 % ) and ro o t s. p o t en t i a l. of. 1 97 1 ) .. the. p la n t. wa s e n h r1 n c e d .. Much o f t h i s c a rbon was ( 5 4 % ) so t h a t t he p r od u c t iv e.

(41) 22 .. The change t o t he r e p r o d u c t ive s t a t e i nv o l ve s maj o r changes in t he d i s t r ib u t i on o f a s s j uli l a t e (vJa rd l aw ,. 1 96 8 ) .. Thi s change i s. d u e ma inly t o t h e i n c re a s i n g s i z e o f t h e s e e d a s a s ink unt i l. even t u a l l y t h e r e q u i reme n t o f t he s e e d d omina t e s a l l o t h e r s i nks s o th a t ac t i v i t y of o t h e r i mp o r t a n t p l a n t p a r t s s lows viz. r o o t s ( Leonard , 1 9 6 2 ;. Ku r s a nov ,. 1 9 6 3 ) vege t a t i ve me r i s t ems ( B ro uwe r ,. Leon ard , 1 9 6 2 ) and n od u l e s ( Lawn a n d B r un , 1 9 7 4 ;. 1 96 2 ;. Have lka , 1 9 7 6 ;. H a r dy and. S i nha , 1 9 7 7 ) .. Thus , a s the p l an t d eve l o ps , t h e t ran s lo ca t ion s t r eam b e comes i n c r ea s in g l y commi t t e d t o s up p l y i ng the s e e d w i th a s s imi l a t e . S o u r c e s o f a s s im i l a t e a re : 1,. Curren t pho t o syn tha t e f rom th e l e a f .. U s ua l l y the pho t os yn t he t i c. area clos e s t t o t he f ru i t p r ov i des t he ma j or p a r t o f thi s supp l y ( F l i nn a n d P a t e , 1 9 7 0 ; 1 97 3 ) .. Love l l a nd Love l l , 1 9 7 0 ;. Hume and C r i swe l l ,. Tran s f e r o f c a rb on a s s i m i l a ted by a f r u i t in g p lan t can b e. t ra n s f e r r e d v e r y e f f i c i en t l y t o t h e s e e d s ( P a te a n d F l in n , 1 9 7 3 ) . Howeve r , Dunp hy and Hanway ( 1 9 7 6 ) sugge s t t h a t the t ran s l o c a t ion o f sugars f rom leaves may b e a l i mi t i n g f a c t o r in s oybe an. p ro duc t ion , 2.. As s imi l a t e s t ored in. t he. l ea f and s tem .. Carbon f i x e d e a r l y i n gro w t h se ems to b e i ne f f i c ient ly t rans fe r r e d t o t h e seed ( Hume a n d C r i s we l l , Probably mos t o f the. 1 97 3 ;. P a t e and F l i nn , 1 9 7 3 ) ,. e a r l y - f i xe d a s s i mi l a t e wh i ch i s r emo b i l i s e d. w a s inc o rp o r a t ed i n p ro t e i ns tha t w e r e hyd r o l i s e d during sene s cence and t r an s f e r and S e t h ,. 1967 ) .. a s amino a c i d s to t h e seeds ( S i mon , 1 9 6 7 ;. Ware ing. Mos t of thi s remo b i l i sa t i on probab l y o c c u r s in. the leave s ( P :i,t e lka ,. 1977 ) .. The 1 0\v e f f i c i ency o f t ran s f e r i s a. consequen c e o f c a rbon b e i n g impo r t a n t i n the s t ru c t u r a l comp onen t s o f the p l an t and b e cause eas i l y mo bi l i se d e a r l y f i xe d carb o n i s d i s s i p a t ed in re s pi r a t i on b e f o r e f l owe r i ng ( P a t e and F l inn , 1 9 7 3 ; 1977) . 3,. Supply f rom the p od , C a r r and P a t e ( 1 9 6 7 ) r e f e r. to. syn th e s i s and t ra n s i e n t s t o r a ge " .. t he pod as "an o r g an o f pho t o I t i s capab le o f p ho t os yn the s i s. ( S inha , 1 9 7 7 ) in c lud ing t ho s e o f l u p i n s ( G r e enwood e t a Z , 3. 1 9 75 ;.

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