leguminosas contenido y vl nutricional.pdf

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E L S E V I E R

Field Crops Research 53 (1997) 3 1-45

F i e l d

C r o p s

R e s e a r c h

L egu m e seeds prote in content and nutr it ional va lue

M a r c e l l o D u r a n t i , C r i s t in a G i u s

Dipartimento di Scienze Molecolari Agroalimentari and Centro lnteruniversitario per lo Studio delle Macromolecole Informazionali

CISMI), Universith di Milano, via Celoria, 2, 1-20133 Milano, Italy

A b s t r a c t

Proteins are ma jor components of legum e seeds. The ir nutritional and functional p roperties dram atically affect the overall

quality of the seed and its technological p erformance. In this review legume seed proteins are considered, taking into account

bot h their mo lecu lar and nutritional properties. Other com pounds of non-protein origin affect the utilization of grain legum es

for food and feed. They hav e also been reviewed both for their involvem ent in the seed l ife cycle and for human nutri t ion.

Th e perspectives f or impro vem ent of legum e traits, starting from the molecules considered and also including the

biotechnological approaches, are also discussed. 1997 Elsevier Science B.V.

1 I n t r o d u c t i o n

G r a i n l e g u m e s a r e w i d e l y r e c o g n i z e d a s i m p o r t a n t

s o u r c e s o f f o o d a n d f e e d p r o t e i n s . In m a n y r e g i o n s

o f t h e w o r l d , l e g u m e s e e d s a r e t h e u n i q u e s u p p l y o f

p r o t e in i n t he d i e t . V e r y o f t e n t he y r e p r e se n t a

n e c e s s a r y s u p p l e m e n t t o o t h e r p r o t e i n s o u r c e s . O n

t h e o t h e r h a n d i n d e v e l o p e d c o u n t r i e s, p l a n t p r o t e i n s

c a n n o w b e r e g a r d e d a s v e r s a t i l e f u n c t i o n a l i n g r e d i -

e n t s o r a s b i o l o g i c a l l y a c t i v e c o m p o n e n t s m o r e t h a n

a s e s se n t i a l nu t r i e n t s . Th i s e vo lu t i on t ow a r ds he a l th

a n d f u n c t i o n a l i t y i s m a i n l y d r i v e n b y t h e d e m a n d s o f

c o n s u m e r s a n d h e a l t h p r o f e s s i o n a l s ( t h e p a r t i a l r e -

p l a c e m e n t o f a n i m a l f o o d s w i t h l e g u m e s i s c l a i m e d

t o i m p r o v e o v e r a l l n u t r i t i o n a l s t a t u s ( G u i l l o n a n d

C h a m p , 1 9 9 6 )) a n d t h e n e e d s o f t h e fo o d i n d u s t r y ,

r e s p e c t i v e l y .

* Corresponding auth or. Dipartimento di Scienze Mo lecolari

Agroalimentari, Univ ersith di M ilano, via Celoria, 2 1-20133

Milano, Italy. Tel.: +39-2-2663 662; fax: + 39-2-70633062.

T h e w o r l d p r o d u c t i o n o f l e g u m e s e e d s , ' t h e p o o r

m a n ' s m e a t ' a s d e v e l o p e d - c o u n tr y p ro d u c e r s c a ll

t h e m , w a s a b o u t 5 8 m i l l i o n t o n s i n 1 9 9 4 ( F A O

e s t im a t ions ) . O f th i s , t he m a jo r pa rt , 40 m i l l i on t ons ,

w a s p r o d u c e d b y d e v e l o p i n g c o u n t r i e s , e s p e c i a l l y

I n d i a a n d C h i n a w i t h o n l y 8 . 5 % c o n s u m e d o u t s i d e

t h e c o u n t r y i n w h i c h i t w a s p r o d u c e d . O n l y A r -

g e n t i n a , M e x i c o , t h e U S A a n d C h i n a e x p o r t s i g n i f i -

c a n t qua n t i t i e s o f l e gum e g r a in , w h i l e Eu r ope i s t he

m a in im por t i ng c on t ine n t ( H e i se r , 1996 ) . P r o t e in s

a r e t h e m a j o r s e e d c o m p o n e n t i n a l l g r a i n l e g u m e s ,

a nd a r e t he r e a son f o r t he i r r e l e va n t nu t r i t i ona l a nd

s o c i o - e c o n o m i c a l i m p a c t . N e w r e s e a r c h a p p r o a c h e s

w h i c h r e l y s t r o n g l y o n b i o t e c h n o l o g y t o i m p r o v e

g r o w t h a n d u t i l i z a t i o n o f g r a i n l e g u m e s h a v e h a d

s ign i f i c a n t im pa c t s on t he nu t r i t i ona l qua l i t y o f

l e g u m e s .

I n t h is r e v i e w w e w i l l f o c u s o n t h e m o l e c u l a r a n d

n u t r i t i o n a l p r o p e r t i e s o f l e g u m e s e e d p r o t e i n s . T h e

p e r s p e c t i v e s f o r t h e i m p r o v e m e n t o f t h e p r o t e in q u a l -

i t y t r a i t s o f g r a in l e gum e s w i l l a r i s e f r om the se

c o n s i d e r a t i o n s . T h e p r o t e i n a n d n o n - p r o t e i n c o m -

0378 -4290 /97/ 17.00 1997 Elsevie r Science B.V . All rights reserved.

PII

S 0 3 7 8 - 4 2 9 0 ( 9 7 ) 0 0 0 2 I - X


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3

M. Duranti, C. Gius / Field Crops Research 53 1997) 31-45

pounds a f f ec t i ng t he ove ra l l nu t r i t i ona l va l ue o f m a-

j o r l egum e c rops wi l l a l s o be cons i de red .

2 T h e s ee d p r o te i n s

L e g u m e s e e d s a c c u m u l a t e l a rg e a m o u n t s o f p r o -

t e i n s d u r i n g t h e i r d e v e l o p m e n t . M o s t a r e d e v o i d o f

ca t a l y t i c ac t i v i t y and p l ay no s t ruc t u ra l ro l e i n t he

c o t y l e d o n a r y t is s u e . T h e y a r e s t o r e d i n m e m b r a n e -

bound o rgane l l e s (p ro t e i n bod i es ) i n t he co t y l e -

d o n a r y p a r e n c h y m a c e l ls , s u r v i v e d e s i c c a ti o n o n s e e d

m a t u r a t i o n a n d u n d e r g o h y d r o l y s i s a t g e r m i n a t i o n ,

t h u s p r o v i d i n g a m m o n i a a n d c a r b o n s k e l e t o n s t o t h e

deve l op i ng s eed l i ngs . S eed p ro t e i n s t ha t behave i n

t h is w a y a r e t e r m e d s t o r a g e p r o t e i n s . H o w e v e r ,

becaus e o f t he i r i n s o l ub i l i t y i n wa t e r and s o l ub i l i t y

i n s a l t s o l u t i ons , t he s t o rage p ro t e i n s a r e a l s o nam ed

g l o b u li n s a n d t h e t w o t e r m s a r e c o m m o n l y u s e d

i n t e rchan geab l y (Derby s h i r e e t a l ., 1976 ) . B es i des

s t o rage p ro t e i n s , l egum e s eeds con t a i n s eve ra l com -

para t i ve l y m i no r p ro t e i n s i nc l ud i ng t ryps i n i n -

h i b i t o r s , l ec t i n s , l i poxygenas e and u reas e , wh i ch a re

re l evan t t o t he nu t r i t i ona l qua l i t y o f t he s eed . S om e

o f t h e m , l i k e u r e a s e f r o m ja c k - b e a n , a l so s e e m t o

h a v e a d o p t e d a s t o ra g e r o l e b y v i r t u e o f t h e i r a m o u n t

i n t he s eed (C as ey e t a l . , 1986 ) .

T h e c a t a l y t i c p r o t e i n s , m o s t o f w h i c h b e l o n g t o

t he vas t bu t l e s s abun dan t c l a s s o f t he a l bum i ns , i . e .

t he p ro t e i n s s o l ub l e i n wa t e r , a r e m a i n l y r ep res en t ed

b y t h e t h o u s a n d s o f d i f f e r e n t e n z y m e s n e c e s s a r y f o r

t h e c o t y l e d o n a r y c el l m e t a b o l i s m a n d w i ll o n l y b e

m arg i na l l y cons i de red i n t h i s r ev i ew.

2 1 The globulins

The g l obu l i n s a r e gene ra l l y c l a s s i f i ed a s 11S and

7S p ro t e i n s acco rd i ng t o t he i r s ed i m en t a t i on coe f f i -

c i en t s . Th e we l l - s tud i ed 11S and 7S p ro t e i n s o f pea

a re nam ed l egum i n and v i c i l i n r e s pec t i ve l y , s o t ha t

t h e c o r r e s p o n d i n g p r o t e i n s o f o t h e r s e e d s a r e o f t e n

re fe r r ed t o a s l egum i n - and v i c i l i n - l i ke g l obu l i n s .

The 7S p ro t e i n s a r e o l i gom er i c p ro t e i n s (u s ua l l y

t r i m e r s ) s h o w i n g p H a n d i o n i c s t r e n g t h - d e p e n d e n t

as s oc i a t i on -d i s s oc i a t i o n equ i l ib r i a . The 1 IS p ro t e i n s

a re a l s o o l i gom ers (u s ua l l y hexam ers ) , bu t a r e l e s s

s us cep t i b l e t o d i s s oc i a t i on , excep t a t ve ry l ow pH o r

i on i c s t r eng t h . Moreover , l a rge r agg rega t es o f 15 -

1 8 S h a v e b e e n r e p o r t e d f o r s o y b e a n l e g u m i n - l i k e

p r o t e in s ( K o s h i y a m a , 1 9 8 3 ). I n t h e p r e s e n c e o f d e n a -

t u r i ng agen t s , s uch as u rea o r s od i um dodecy l s u l -

pha t e , bo t h the l l S and 7S p ro t e i n s li be ra t e t he i r

cons t i t uen t po l ypep t i de cha i n s . Thes e po l ypep t i des

a r e n a t u r a l l y h e t e r o g e n e o u s : t h o s e w h i c h h a v e b e e n

p u r i f i e d t o h o m o g e n e i t y w i l l o f t e n a p p e a r t o b e

m i x t u r e s o f d i f f e r e n t m o l e c u l a r s p e c i e s, i f e x a m i n e d

by o t he r m e t hods (P us z t a i and S t ewar t , 1980) . He t -

e rogene i t y i s ev i den t a t bo t h s i ze and cha rge l eve l s

(B ro wn e t a l. , 1981 ; Tuc c i e t a l ., 1991 ; Ho rs t m an n e t

a l. , 1993 ) , and a r i s es f rom a com bi na t i on o f t wo

fac t o r s , t he m u l t i gene o r i g i n o f each s t o rage g l obu l i n

(C as ey e t a l . , 1986 ) and t he pos t - t r ans l a t i ona l m od i -

f i c a t i o n s o f r e l a t i v e l y f e w e x p r e s s i o n p r o d u c t s

(W r i gh t , 1986 ) . The r e l a t i ve con t r i bu t i on o f t hes e

fac t o r s va ry s i gn i f i can t ly i n t ra - and i n t e r -gene r i ca l l y .

I t appea r s t ha t t he ru le s fo r t he a s s em b l y o f na t i ve

l l S and 7S p ro t e i n s f rom t he i r he t e roge neou s s ub -

un i t s a r e no t we l l de f i ned and t ha t r andom as s oc i a -

t i on m i gh t oc cu r i n m an y i n s t ances (Ga t eh ous e e t a l .,

1981 ; Akazawa and Hara -Ni s h i m ura , 1985 ) , t hus

e x p o n e n t i a l l y i n c r e a s i n g t h e n u m b e r o f m o l e c u l a r

s pec i es i n each c l a s s. Th e s i tua t i on i s fu r t he r com pl i -

ca t ed by t he f ac t t ha t m any o f t hes e p ro t e i n s ( e s pe -

c i a l l y 7S ) a re uneven l y g l ycos y l a t ed .

He t e roge ne i t y i s t he gene t i c and m o l ecu l a r s t a rt -

i n g p o i n t f o r m o d e m D N A - r e c o m b i n a n t t e c h n o l o g i e s

a i m e d t o i m p r o v e t h e o v e r a l l p e r f o r m a n c e o f l e g u m e

prote ins .

2 1 1 The l l S globulins

T h e w i d e l y a c c e p t e d s t ru c t u ra l m o d e l o f 1 1S p r o -

t e i n s i s s t i l l t ha t p ropos ed fo r Vicia faba l e g u m i n

(P l i e t z e t a l. , 1984 ) . In t h i s m ode l s i x m o nom ers a re

a r r anged i n a t r i gona l an t i p r i s m com pac t s t ruc t u re ,

w h i c h i s n e c e s s a r y to t h e d e n s e p a c k i n g o f th e

m o l e c u l e s i n t h e w a t e r - p o o r m e d i u m o f t h e p r o t e i n

b o d i e s. T h e m o n o m e r i c u n i t o f v i r t u a ll y a l l l e g um i n -

l i ke g l obu l i n s cons i s ts o f an ac i d i c po l ype p t i de cha i n

( r ang i ng i n s i ze f rom 25 ,000 t o 50 ,000 da l t ons (C as ey

e t a l . , 1986 ) ) d i s u l ph i de -bonded t o a bas i c cha i n

(usual ly around 20 ,000) , a l so named c~ and 13 sub-

uni t s respec t ively ; the c~ and /3 subuni t s are equim o-

l a r , s i nce , a s we wi l l s ee be l ow, t hey a r i s e f rom t he

pos t - t r ans la t i ona l p ro t eo l y t i c c l eavag e o f an c~ + [3

p r e c u r s o r p o l y p e p t id e .


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M. Duranti, C. Gius Field Crops Research 53 1997) 31-45 33

T h e a + / 3 p o l y p e p t i d e s o f l e g u m i n - l ik e p r o t e in s

a re s yn t hes i zed a s p re -p ro -po l ypep t i des . The f i r s t

p ro t eo l y t i c even t cons i s t s i n t he co - t r ans l a t iona l r e -

m o v a l o f t h e h i g h l y h y d r o p h o b i c s i g n a l p e p t i d e

(B l obe l and Dobber s t e i n , 1975 ; Mi i n t z e t a l . , 1985 ) ,

w h i c h p r o d u c e s a p r o - p o l y p e p t i d e , w h e r e f o r m a t i o n

o f i n t r am o l ecu l a r d i s u l ph i de b r i dge ( s ) t akes p l ace . A

fu r t he r l i m i t ed p ro t eo l y t i c deg rada t i on , oc cu r r i ng i m -

m ed i a t e l y a f t e r t he depos i t i on o f t he l egum i n i n t he

p r o t e in b o d i e s, g i v e s ri s e to t h e - S - S - l i n k e d c~ a n d

/ 3 m a t u re cha i n s (Ga t ehous e e t a l . , 1984 ; Mi i n t z ,

1 9 8 9) . C l e a v a g e o f t h e p r o - p o l y p e p t i d e o c c u r s b e -

t w e e n a n A s n - G l y p e p ti d e b o n d in p r o x i m i ty o f a

h i g h l y h y d r o p h y l i c r e g i o n (N i e l s en , 1 9 8 4) . A c o n -

v e r t a s e w h i c h h a s b e e n i s o l a t e d a n d c h a r a c t e r i z e d i s

r e s pons i b l e fo r t h i s s pec i f i c c l eavage (S co t t e t a l . ,

1 9 9 2 ; M u r u m a t s u a n d F u k a z a w a , 1 9 9 3 ; S h i m a d a e t

a l ., 1994 ) . Th e p ro t eo l y t i c m o d i f i ca t i on o f legum i n

p ro -p o l ype p t i de t o m a t u re a and /3 cha i n s tr i gge r s

t h e c o n v e r s i o n o f t h e t r i m e r i c p r o - p o l y p e p t i d e a s -

s e m b l y t o t h e h e x a m e r i c a s s o c i a ti o n o f t h e m a t u r e

legumin (Niel sen e t a l . , 1989; Durant i e t a l . , 1992) .

T h e m o l e c u l a r b a s i s o f t h i s p h e n o m e n o n w i l l b e

m a d e c l e a r o n c e t h e c r y s t a ls o f b o t h l e g u m i n f o r m s

wi l l be ava i l ab l e fo r X- ray ana l y s i s .

A s i ng l e po i n t m u t a t i on o f lup i n 11S g l obu l i n has

rea l i zed a g l yc os y l a t i on s it e , by v i r t ue o f wh i ch l up i n

1 I S g l o b u li n i s t h e o n l y k n o w n g l y c o s y l a t e d p r o te i n

of th i s c lass (Dura nt i e t a l ., 1995) .

2.1.2. The 7S globulins

The 7S g l obu l i n s o r v i c i l i n - l i ke p ro t e i n s a r e even

m o r e h e t e r o g e n e o u s t h a n l e g u m i n - l i k e p r o t e i n s , d u e

t o a n u n e v e n g l y c o s y l a t i o n o f t h e i r s u b u n it s ( S c h o l z

e t a l. , 1983 ) . As an exam pl e , phas eo l i n can ex i s t i n a

c o m b i n a t i o n o f d i f f e r e n t i a l l y g l y c o s y l a t e d p o l y -

pept ides (Bol l in i e t a l . , 1983) .

Th e gen era l l y -ac cep t e d s t ruc t u re fo r v i c i li n is a

t r im e r o f p o l y p e p ti d e s o f M r 4 0 , 0 0 0 - 7 5 , 0 0 0 m a k i n g

u p a n a t iv e p r o te i n o f M r a b o u t 1 5 0 , 0 0 0 - 1 7 0 , 0 0 0

(P l i e t z e t a l . , 1983 ) . The v i c i l i n s no rm al l y do no t

c o n t a i n c y s t e i n e s a n d t h e r e f o r e h a v e n o d i s u l p h i d e

bonds (C as ey e t a l . , 1986 ) .

M o s t v i c i l i n s u n d e r g o a m o r e e x t e n s i v e p r o t e -

o l y t i c p roces s i ng t han t he l egum i ns , r e s u l t i ng i n a

n u m b e r o f p o l y p e p t i d e fr a g m e n t s w h i c h , d e s p i te t h e

l o s s o f cova l en t con t i nu i t y , s t i l l k eep t oge t he r i n an

as s oc i a t ed s t ruc t u re (Ga t ehous e e t a l . , 1983 ; Mon-

s a l ve e t a l. , 1990 ) . Th e s ch em e fo r de r i va t i on o f t he

p o l y p e p t i d e f r a g m e n t s f r o m t h e 5 0 , 0 0 0 p r o - p o l y -

p e p t i d e in p e a h a s b e e n m a d e p o s s i b l e b y c o m p a r i n g

a m i n o a c i d s w i t h c D N A s e q u e n c e s ( G a t e h o u s e e t a l. ,

1983; Lyc et t e t a l . , 1983; Spe nce r e t a l ., 1 983) .

B i os yn t he t i c s t ud i es have i nd i ca t ed t ha t Pisum vi -

c i l i n p recu r s o r po l ypep t i des , l i ke t hos e o f t he l egu -

m i n p recu r s o r s , a r e s yn t hes i zed wi t h a N- t e rm i na l

s i gna l pep t i de (H i gg i n s and S pence r , 1981 ) , wh i ch i s

co - t r ans l a t i ona l l y r em oved i n s i de t he endop l as m i c

re t i cu l um . O ne o f the m os t s t ud i ed v i c i l in s i s phas e -

o l i n , t he m a j o r s eed s t o rage p ro t e i n f rom Phaseolus

vulgaris (B o l l i n i and C hr i s pee l s , 1978 ) . I t s t h ree -d i -

m e n s i o n a l s t r u c t u r e h a s b e e n d e t e r m i n e d ( L a w r e n c e

et a l. , 1990; K o et a l. , 1993; L aw ren ce e t al ., 1994) ,

t hus p rov i d i ng a power fu l t oo l fo r t he i nves t i ga t i on

o f t h e s t r u c t u r e - f u n c t i o n r e l a t i o n s h i p s a n d f o r t h e

m an i pu l a t i on o f i t s nu t r i t i ona l and func t i ona l p rope r -

t i e s (Dyer e t a l . , 1993 ) . P has eo l i n i s an he t e roge -

n e o u s t r i m e r o f M r b e t w e e n 1 4 0 ,0 0 0 a n d 1 6 0 ,0 0 0

(S un e t a l ., 1975 ; B o l l i n i and C hr i s pee l s , 1978 ) . Th e

t h ree s i ze c l a s s es o f phas eo l i n po l ype p t i des a re s t ruc -

t u ra l l y s i m i l a r and appea r t o be encoded by a r e l a -

t i v e ly s m a l l a n d c o n s e r v e d m u l t ig e n e f a m i l y ( T a l b o t

e t a l. , 1984 ) . P has eo l i n i s a g l ycop ro t e i n w i t h t wo

po t en t i a l g l ycos y l a t i on s i te s (S l i gh t om e t a l. , 1983 ) .

D i f f e ren t i a l g l ycos y l a t i on can occu r a t e i t he r s i t e

g i v in g r i s e t o f u r t h e r p o l y p e p t i d e h e t e r o g e n e i t y ( L i o i

and Bol l in i , 1983) .

D e s p i t e t h e h e t e r o g e n e i t y o f s t o r a g e p r o t e i n s

am ong s pec i es and even wi t h i n a s i ng l e s pec i es

(Ni e l s en e t a l ., 1989 ) , l l S and 7S g l obu l i n s hav e

a l s o been i den t i f i ed i n ce rea l s (Wen and Lu t he ,

1985 ) and non -ce rea l s eeds (S i m on e t a l . , 1985 )

s ugges t i ng t ha t t he genes i nvo l ved a re anc i en t and

o r i g i na t ed l ong be fo re s pec i a t i on o f t he ances t r a l

fo rm s o f t he m an y d i ve r s e t axa (C as ey e t a l ., 1986 ) .

S u p p o s i t io n o f a c o m m o n a n c e s t ra l o r i g in o f 7 S a n d

11S s eed s t o rage p ro t e i n s i s con f i rm ed on a m o l ecu -

l a r and gene t i c bas i s (B o r ro t o and Dure I I I , 1987 ;

Gibbs e t a l . , 1989; Shutov e t a l . , 1995) ; despi te

ex t r em e d i ve r s i t y o f t hes e f am i l i e s o f s eed g l obu l i n s ,

t h e r e s e e m s t o b e h i g h a m i n o a c i d s e q u e n c e c o n s e r -

va t i on and s t ruc t u ra l r e l a t i ons h i p am ong t hem (Argos

e t a l . , 1985 ) . F rom t h i s po i n t o f v i ew t he g l obu l i n s

c a n a l s o b e r e g a r d e d a s e v o l u t i o n a r y m a r k e r s o f th e

d i f f e ren t l egum e s pec i es .


4/15

34

M. Duranti, C. Gius / Field Crops Research 53 1997) 31-4 5

3 Nutr i t iona l va lue o f l egume s eed prote ins

A l l l egume s eed p ro te in s a r e r e l a t ive ly low in

s u lphu r - con ta in ing amino ac id s and t ryp tophan , bu t

the amoun t s o f ano the r e s s en t i a l amino ac id , ly s ine ,

a r e mu ch g rea te r than in ce r ea l g ra in s (Rock land and

Radke, 1981; Ampe et a l . , 1986) . Therefore , wi th

r es pec t to ly s ine and s u lphu r amino ac id con ten t s ,

ce r ea l and l eg um e p ro te in s a r e nu t r i t iona l ly comple -

m e n t a r y . T h e d e g r e e o f m u t u a l s u p p l e m e n ta t i o n m a y

a l s o depend , how ever , on the con ten t s o f s econd

l imi t ing amino ac id s , i . e . t h r eon ine in ce r ea l s and

t ryp tophan in l egumes .

The es s en t i a l and no n -es s en t i a l amino ac id pa t t e rn

o f e l even ma jo r l egume s eeds , inc lud ing up to 3

species in each genus , i s shown in F ig . 1 .

H ow ever , amino ac id compos i t ion r ep res en t s the

po ten t i a l qua l i ty o f a p ro te in food , the i r b ioava il ab i l-

i ty be ing c r i t ica l f o r the s upp ly o f amino ac id s in the

d ie t . A number o f exper imen ta l app roaches , dev i s ed

to a s s es s the b ioava i l ab i l ity o f amino ac id s in l egum e

foods , nam ely P ro te in Ef f i c i enc y Ra t io (P ER) , N e t

P ro te in Ra t io (N P R) , Re la t ive N P R (RN P R) (S a rw ar

and M cD on ough , 1990), and True P ro te in D iges t ib i l -

i t y -c o r r e c te d A m i n o A c i d S c o r e ( A A S DTP

( F A O / W H O ( 19 9 0 )) , c o n c u r r e n tl y d e m o n s t r at e t h a t

s eed p ro te in s have a low er ove ra l l nu t r i t iona l qua l i ty

than an im a l p ro te in s . Th i s can be r e l a t ed to the i r low

con ten t o f s u lphu r - con ta in ing amino ac id s (S a rw ar

and Peace, 1986) , the compact pro teolys is - res is tant

s t ruc tu re o f na t ive s eed p ro te in s (D es phande and

Nielsen , 1987) and the presence of ant inutr i t ional

Arg I

i l e i i

L I

n | i

Val I 5 ,

T h r i ~ i

M e t ~ 4 . 0

e ~ ~ 4 o

~

I

~ I I I I

c ~ i 1 ~ 1

S e r I l

A s x

I

t a x i

o |

m i

e i

E A A

8 0 1 2 . 0 1 6 . 0 2 4 . 0

0 . 0 1 2 . 0 1 6 . 0 2 4 . 0

l

N E A A

F i g . 1 . A m i n o a c i d c o n t e n t o f s e l e c te d l e g u m e s . T h e l o w e s t w h i t e b a r s ) a n d t h e h i g h e s t b l a c k b a r s ) c o n t e n t o f e a c h a m i n o a c i d a m o n g t h e

f o l l o w i n g l e g u m e s e e d s i s g i v e n : a :

Cajanus cajan

Ne me e t a l . , 1975 ), b :

Canavalia ensiformis

Mabesa, 1984), c :

Cicer arietinum

A t t i a e t

al. , 1994), d:

Glycine max

Wang and Chang , 1995 ) , e :

Lens culinaris

Hsu et a l . , 1980), f :

Lupinus albus

Chango e t a l . , 1995), g :

Lupinus

angustifolius

Hove, 1974), b :

Lupinus luteus

Chango e t a l . , 1995), i :

Pisum sativum

Le term e et a l . , 1990), l :

Phaseolus lunatus

All i e t

a l . , 1994), m:

Phaseolus mungo

M abesa , 1984 ) , n :

Phaseolus vulgaris

Steel e t a l . , 1995), o :

Vicia faba

Hebb le thwi te , 1983 ) , p :

Vicia

angustifolia

M abesa , 1984 ) , q :

Vigna unguiculata Khan

e t a l ., 1 9 80 ). V a l u e s a r e g i v e n a s g / 1 0 0 g p r o t e i n . E A A : e s s e n t i a l a m i n o a c i d s .

N E A A : n o n - e s s e n t i a l a m i n o a c i d s .


5/15

M. Duranti , 6 . Gius / F ie ld Crops Research 53 (1997) 3 1- 45

5

c o m p o u n d s , w h i c h m a y a f f e c t d ig e s t ib i l it y o f p r o -

t e in s t h e m s e l v e s a n d o t h e r c o m p o n e n t s L i e n e r , 1 9 7 9;

B res s an i e t a l . , 1982 ; Aw and S wans on , 1985 ) .

4 Legume seed ANtinutr itional C om poun ds

A N C s )

Legum e s eeds con t a i n s eve ra l an t i nu t r i t i ona l p ro -

t e i n a n d n o n - p r o t e i n c o m p o u n d s . T h e p r e s e n c e o f

ANC s i n c rop p l an t s i s o f t en t he r e s u l t o f an evo l u -

t i ona ry adap t a t i on wh i ch enab l es t he p l an t t o s u rv i ve

and com pl e t e i t s l i f e cyc l e under na t u ra l cond i t i ons ,

r e g a r d l e ss o f t h e n e g a t i v e c o n s e q u e n c e s o n t h e q u a l -

i t y a n d s a f e t y o f t h e f o o d p r o d u c t s . I n d e e d , d u e t o

t he i r an t i nu t r i t i ona l o r even t ox i c p roper t i e s , va r i ous

p o t e n t i a ll y h a r m f u l c o m p o u n d s h a v e b e e n s h o w n t o

p l a y a p r o t e c ti v e r o l e a g a in s t in s e c t s P e u m a n s a n d

van Dam m e , 1995 ) , fung i J e l l is and Vas s i e , 1995 ),

p r e d a t o r s a n d a n u m b e r o f s t r e s s c o n d i t i o n s

C hr i s pee l s and R a i khe l , 1991 ) .

4 1 Protein ANCs

S e e d h y d r o l a s e i n h i b i t o r s a r e c o n s i d e r e d t o b e

i m p o r t a n t i n d e t e r m i n i n g t h e q u a l i ty o f l e g u m e s e e d s

and a re c l a s s i f i ed a s an t i nu t r i t i ona l com pounds . The i r

an t i nu t r i t i ona l e f f ec t i n t he i r r eve r s i b l e i nh i b i t i on o f

v a r i o u s d i g e s t i v e e n z y m e s i s w e l l d o c u m e n t e d

Le term e et a l. , 1992) .

H o w e v e r , t h e i r e f f e c t i s u s u a l l y m a n i f e s t o n l y i f

t h e s e e d o r t h e f l o u r a r e c o n s u m e d u n c o o k e d , s i n c e

hea t dena t u ra t i on no rm al l y i nac t i va t e s t hes e p ro t e i n s

Vi da l -Va l ve rde e t a l . , 1994 ) . Once i nac t i va t ed , t he

p ro t e i n i nh i b i t o r s m ay even p l ay a pos i t i ve nu t r i -

t i ona l ro l e , due t o t he i r h i gh con t en t o f s u l phu r -con -

t a i n i ng am i no ac i d s r e l a t i ve t o t he m a j o r i t y o f t he

s e e d p r o t e in s R y a n , 1 9 9 0) . T h e m o s t c h a r a c t e r i z e d

p ro t e i n inh i b i t o r s a re T ryps i n Inh i b i to r s T I s ) o f bo t h

t h e B o w m a n - B i r k t y p e D o m o n e y et a l., 1 9 93 ) i n

Pisum sativum a n d K u n i t z t y p e i n Glycine max

H o r i s b e r g e r a n d T a c c h i n i - V o n l a n t h e n , 1 9 8 3 ) a n d

a - a m y l a s e i n h ib i to r s M o r e n o e t a l. , 1 9 9 0 ). R e c e n t l y

an i n v i t ro d i ges t i on s y s t em has been u s ed t o eva l u -

a t e t h e i m p o r t a n c e o f p e a s e e d T I i n p r o t e in d i g e s -

t i on Do m one y e t a l ., 1993 ) . A r e l a t i ve l y l ow l eve l

o f i n h i b i t i o n i s o b s e r v e d w h e n p u r i f i e d p e a T I s a r e

e v a l u a t e d u s i n g t h i s s y s t e m , c o m p a r e d t o e x p e r i -

m e n t s w h e r e s o y b e a n T I s a r e u se d . M o r e o v e r , v a r ia -

t i on i n pea p ro t e i n d i ges t i b i l i t y has been obs e rved

a m o n g a s e t o f p e a l in e s t h a t a r e n e a r - is o g e n i c a n d

con t a i n s i m i l a r s eed t ryps i n i nh i b i t o r ac t i v i t i e s , s ug -

ges t i ng t ha t s eed com pounds , o t he r t han i nh i b i t o r s ,

m a y b e o f g r e a t e r i m p o r t a n c e i n a f f e c t i n g p r o t e i n

diges t ib i l i ty A1-W esal i e t a l ., 1995) .

T I s h a v e b e e n p r o v e n t o a c t a s p r o t e c t i v e a g e n t s

aga i n s t in s ec t a t tack Hi l de r e t al. , 1990 ; J ohns on e t

a l ., 1989 ) and t he i r i nvo l vem en t i n woun d r e s pons es

i s cu r r en t l y r ece i v i ng a t t en t i on in a nu m b er o f s pec i es

Gros jean e t a l . , 1993) .

S eed l ec t i n s a r e s uga r -b i nd i ng p ro t e i n s , a s we l l a s

hem agg l u t i n i n s , wh i ch a re ab l e t o agg l u t i na t e r ed

b l ood ce l l s E t z l e r , 1985 ) . M any bu t no t a ll , lec t i n s

have hem agg l u t i na t i ng ac t i v i t y L i s and S haron ,

1986 ) . They a re found i n m os t p l an t p roduc t s i nc l ud -

i n g t h o s e t h a t m a y b e e a t e n w i t h o u t h e a t - tr e a t m e n t o r

p roces s i ng l i ke Solanaceae S haron and L i s , 1990 ) .

T h e t o x i c i t y o f l e c t in s i s c h a r a c t e r i z e d b y g r o w t h

i nh i b i t i on i n exper i m en t a l an i m a l s , and by d i a r rhea ,

n a u s e a , b l o a t i n g a n d v o m i t i n g w h e n i n j e c t e d i n h u -

m ans . S om e l ec t i n s can caus e agg l u t i na t i on o f t he

r e d b l o o d c e l l s f o l l o w e d b y h e m o l y s i s a n d , i n e x -

t r em e cas es , dea t h L i en e r e t a l ., 1986 ). H ea t p ro -

ces s i ng can r educe t he t ox i c i t y o f l ec t i n s , bu t l ow

t e m p e r a t u r e o r i n s u f f i c i e n t c o o k i n g m a y n o t c o m -

p l e t e l y e l i m i na t e t he i r t ox i c i t y van Dr i es s che , 1988 ) .

A l t hou gh t he ro l e o f l ec ti n i n l egum e s eeds i s s ti ll

con t rove r s i a l , t he re i s ev i dence t ha t l ec t i n s a r e de -

f ens e p ro t e i n s aga i n s t po t en t i a l p l an t enem i es . S i nce

t he m a j o r i t y o f p l an t l ec t i n s exh i b i t a s pec i f i c i t y

aga i n s t ca rbohyd ra t e s o f an i m a l o r i g i n , one can r ea -

s onab l y a rgu e t ha t p l an t s u s e t h i s t ype o f p ro t e i n a s a

d e f e n s e a g a i n s t p h y t o p h a g o u s i n v e r t e b r a te s a n d h e r -

b i vo rou s an i m a l s . F eed i ng t r i a l s con f i rm t he de l e t e r i -

ous e f f ec t o f va r i ous p l an t l ec t i n s on i n s ec t s and r a t s .

However due t o t he i r l i m i t ed t ox i c i t y , l ec t i n s p roba-

b l y d i d no t ev o l ve a s p ro t ec t an t s o f i nd i v i dua l p lan t s ,

bu t r a t he r a s r e s i s t ance f ac t o r s wh i ch a re bene f i c i a l

f o r t h e s u r v iv a l o f th e s p e c i e s P e u m a n s a n d v a n

D a m m e , 1 9 9 5 ) .

A l l e rg i e s t o l egum e s eeds a re r e l a t i ve l y uncom -

m o n i n h u m a n s d u e t o t h e l o w a l l e r g e n ic c a p a c i ty o f

s t o rage p ro t e i n s La l l e s and P e lt r e , 1996 ) , bu t t hey

c o u l d d e v e l o p w i t h i n c r e a s e d c o n s u m p t i o n , a s e x e m -


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6 M. Duranti, C. Gius / FieM Crops Research 53 1997) 31-45

p l i f i e d b y p e a n u t a n d s o y b e a n , w h o s e m a j o r a l l e r -

gens a re w e l l - cha rac t e r i zed p ro t e i n s . In t e res t ing ex -

pe r i m en t s on i n t e s t i na l s u rv i va l , ab s o rp t i on and an t i -

gen i c i t y o f s oybean an t i nu t r i en t s i n t he r a t (Gyongy i

e f a l. , 1995 ) s how ed t ha t t he e l i m i na t i on o r s ubs t an -

t i a l r educ t i on o f t h i s undes i r ab l e b i o l og i ca l ac t i v i t y

wou l d s ubs t an t i a l l y i nc reas e t he nu t r i t i ve va l ue o f

s o y b e a n p r o d u c t s . M o r e o v e r , s t u d i e s o f p r o t e o l y t i c

m o d i f i c a t i o n a n d m e t h i o n i n e e n r i c h m e n t o f s o y p r o -

t e i n an t i gens s howed a r educ t i on i n a l l e rgen i c i t y and

a n o v e r a l l i m p r o v e m e n t o f t h e s o y b e a n m e a l v a l u e

(Hajos e t a l . , 1993) .

4 . 2. N o n - p r o t e i n A N C s

L e g u m e s e e d s c o n ta i n a n u m b e r o f n o n - p r o t e i n

ANC s wi t h s i gn i f i can t l y d i f f e ren t s t ruc t u res and e f -

f e c t s . T h e m o s t i m p o r t a n t o f t h e m a r e l i s t e d b e l o w .

A l k a l o i d s l i m i t th e a c c e p t a n c e o f v a r i o u s l e g u m e

s eeds , s uch as l up i n , bo t h fo r t he i r s t rong b i t t e r ta s t e

and t ox i c i t y (M ark i ev i cz e t a l. , 1988 ; C uad ra do e t

a l ., 1992 ) . Th e b i t t e r s eeds con t a i n up to 2 a l ka -

l o i d s , u s ua l l y l up i n i ne , cy t i s i ne , s pa r t e i ne , l upan i ne

and anagy r i ne . Due t o t he wa t e r s o l ub i l i t y o f a l ka -

l o i d s and t he i r l ow s i ze , i t i s pos s i b l e t o r em ove t he m

f r o m t h e s e e d s b y s o a k i n g a n d c o o k i n g i n w a t e r

(S ch oen eber ge r e t a l ., 1982 ) . Lup i n va r i e t ie s w i t h

l ow a l ka l o i d s (0 .04 ) , wh i ch a re s u i t ab le to hum an

a n d a n i m a l c o n s u m p t i o n w i t h o u t d e b i t t e r i n g

(C ua d rado e t a l ., 1992 ) , have bee n s e l ec t ed (Har r i s on

and Wi l l i am s , 1982 ) . A p robab l e ro l e o f a l ka l o i d s

can a l s o be t o p rov i de t he s eeds pes t p ro t ec t i on . In

add i t i on , a l ka l o i d s ex t r ac t ed f rom l up i n s eeds can be

u s e d f o r p h a r m a c o l o g i c a l a n d o t h e r b i o m e d i c a l p u r -

pos es (De l a C uad ra e t a l . , 1992 ) .

T o g e t h e r w i t h a l k a l o i d s , o t h e r c o m p o u n d s m a y

con t r i bu t e t o t he s ca rce pa l a t ab i l i t y o f s om e l egum e

s eeds . Th i s i s t he cas e o f Vi c i a nar bonens i s s eeds

w h e r e t h e p r e s e n c e o f y - g l u t a m y l - e t h e n y l - c y s t e i n e

c a u s e s t h e f o r m a t i o n o f s u l p h u r o u s c o m p o u n d s o n

e n z y m a t i c h y d r o l y s i s ( P e t e r E i c h i n g e r , p e r s o n a l

c o m m u n i c a t i o n ) .

P hy t i c ac i d , m yo i nos i t o l 1 , 2 ,3 ,4 ,5 ,6 hexak i s -d i hy -

d rogen phos pha t e , i s p res en t i n l egum e s eeds , m ak -

i ng up t he m a j o r po r t i on o f t he t o t a l phos pho rous i n

t he s eed . P hy t i c ac i d i s r e s pons i b l e fo r t he r educ t i on

o f t he b i oava i l ab i l i t y o f e s s en t i a l m i ne ra l s , fo rm i ng

i n s o l ub l e com pl exes wh i ch a re l e s s ava i l ab l e fo r

d i ges t i on and abs o rp t i on i n t he s m a l l i n t e s t i ne (De-

s p h a n d e a n d C h e r y a n , 1 9 8 4 ) . M o r e o v e r , p h y t a t e s

have a l s o been s hown t o i nh i b i t t he ac t i v i t y o f

s eve ra l enzym es (Knuck l es e t a l . , 1989 ) .

P heno l i c com pounds , s uch as t ann i n s , can c ro s s -

l i nk wi t h p ro t e i n s by r eac t i ng wi t h l y s i ne o r m e t h i on -

i ne r e s i dues , m ak i ng t hem unava i l ab l e du r i ng d i ges -

t i on (Dav i s , 1981 ) . J a f f6 (1950 ) had a l r eady s ug -

ges t ed t ha t t he re m ay be an i nve r s e r e l a t i ons h i p

be t ween p ro t e i n d i ges t i b i l i t y and s eed -coa t co l o r i n

Phas eo l u s vu l gar i s whi ch con t a i n t ann i n s and o t he r

p o l y p h e n o l s .

S a p o n i n s a r e a d i v e r s e g r o u p o f c o m p o u n d s c o m -

m o n l y f o u n d i n l e g u m e s ( H u d s o n a n d E 1 - D i f r a w i ,

1979 ) . The i r gene ra l s t ruc t u re cons i st s o f a s t e ro i d o r

t r i t e rpene g roup l i nked t o one o r m ore s uga r

m o l e c u l e s . T h e p r e s e n c e o f b o t h p o l a r a n d n o n - p o l a r

g roups p rov i de s apon i ns w i t h s t rong s u r f ace -ac t i ve

p roper t i e s wh i ch a re r e s pons i b l e fo r t he i r adve r s e

b i o l o g i c a l e f f e c t s . A w e l l k n o w n t o x i c e f f e c t o f

s apon i ns i s t he i r ab i l i t y t o l y s e e ry t h rocy t es , a s we l l

a s o t he r ce l l s , s uch as t hos e found i n t he i n t e s t i na l

m ucos e , t hus a f f ec t i ng nu t r i en t ab s o rp t i on (Huds on ,

1979) .

V i c i n e a n d c o n v i c i n e a r e c o n f i n e d t o Vicia a n d

a r e k n o w n t o b e r e s p o n s i b l e f o r a t y p e o f h e m o l y t i c

a n e m i a , k n o w n a s f a v i s m ( H u d s o n , 1 9 7 9) .

L e g u m e s a r e w e l l k n o w n i n d u c e r s o f i n t e s t i n a l

gas s es ( f l a t u l ence ) , due t o t he p res ence o f a -D-

g a l a c t o p y r a n o s y l r e s id u e s b o u n d t o t h e g l u c o s e m o i -

e t y o f s uc ros e . An i m al s and m an a re no t ab l e t o

d i ges t s uch o l i gos acchar i des , becaus e o f t he abs ence

o f a -ga l ac t o s i das e i n t he i r i n t e s t i na l m ucos e . C ons e -

quen t l y t he a -ga l ac t o s i des pas s i n t o t he co l on and

are f e rm en t ed by t he i n t e s t i na l bac t e r i a w i t h p roduc-

t i on o f gas (F l em i ng , 1981 ) . It i s c l ea r l y des i r ab l e t o

d e c r e a s e t h e o l i g o s a c c h a r i d e c o n t e n t o f l e g u m e s , i f

t hey a re t o be m o re e f f ec t i ve l y exp l o i t ed a s r e l a t i ve l y

i nexp ens i ve s ou rces o f p ro t e i n (Gda l a e t a l. , 1995 ) .

M a n y l e g u m e s h a v e s t a r c h y s e e d s . E v e n i f m o s t

o f t he s t a r ch i s d i ges t ed i n t he upper pa r t o f t he

d i ges t i ve t r ac t , s om e r es i s t an t s t a r ch , u s ua l l y encap -

s u l a t ed i n t he ce l l wa l l s and accoun t i ng fo r up t o

20 o f t he t o t a l s t a r ch (Noa h e t a l. , 1995 ), m a y

e s c a p e d i g e s t io n b y e n d o g e n o u s e n z y m e s o f t h e g a s -

t ro in tes t inal t ract , thus cont r ibut ing to in tes t inal fer -

m en t a t i on .


7/15

M. Duranti, C. Gius / Field Crops Research 53 1997) 31-45 37

5 E f f e ct o f p r o c e s s in g a n d g e r m i n a t i o n o n t h e

n u t r i t i o n a l a n d t e c h n o l o g i c a l q u a l i t y o f l e g u m e

s e e d s

R e m o v a l o f u n d e s i r a b le c o m p o n e n t s i s e s s e n ti a l

t o i m p r o v e t h e n u t r i t i o n a l q u a l i t y o f l e g u m e s a n d

ef fec t i ve l y u ti l ize t he i r fu l l po t en t i a l. R educ t i on i n

t h e a m o u n t s o f m o s t A N C s , i n c l u d in g l e c ti n s R e d d y

and P ea r s on , 1994 ) and ga l ac t o - s ugar s Vi da l -

Va l ve rde e t a l . , 1992 ) , e i t he r by t echno l og i ca l and

h o u s e p r o c e s s i n g o r b y e n d o g e n o u s e n z y m a t i c c a t a l -

y s i s d u r i n g s e e d g e r m i n a t i o n h a v e b e e n r e p o r t e d .

D e c r e a s e o f A N C s m a y o c c u r e i th e r b y p h y s ic a l

r em ova l o r by hea t i nac t i va t i on , e s pec i a l l y wi t h p ro -

t e i n ANC s . C ook i ng genera l l y i nac t i va t e s hea t - s ens i -

t i ve f ac t o r s s uch as p ro t e i n i nh i b i t o r s Tab l e 1 ) . In

m a n y i n s ta n c e s , th e u s e o f o n l y o n e m e t h o d m a y n o t

c o m p l e t e l y r e m o v e a g i v e n A N C a n d a c o m b i n a t i o n

o f t w o o r m o r e m e t h o d s is r e q u i r e d V i d a l - V a l v e r d e

e t a l. , 1992 ) . P o l yph eno l s a re s i gn i f i can t l y r edu ced

b y s o a k in g a n d c o o k i n g . O n t h e o t h e r h a n d , t h e

con t en t o f t ann i n s and ca t ech i n s i n l en t i l s and b road

beans s eem s t o i nc reas e a f t e r t hes e t r ea t m en t s

Vi da l -Va l ve rde e t a l . , 1994 ) . Th i s can be a t t r i bu t ed

t o a g rea t e r ac ces s i b i l i ty t o ana l y s i s o f t ann i n s , wh i ch

a re o r i g i na l l y i n t he i n t e r i o r o f t he ce l l and o f t en

l in k e d to p r o te i n s a n d o t h e r m a c r o m o l e c u l e s H a g e r -

m an , 1992 ) .

S e e d g e r m i n a t io n a l s o h a s a d o c u m e n t e d e f f e c t in

t he r em ova l o f an t i nu t r i en t s i n l egum es , s i nce i t

m o b i l i z e s t h o s e c o m p o u n d s w h i c h a r e t h o u g h t t o

func t i on a s r e s e rve nu t r i en t s , e . g , phy t a t e s , o l i go -

s acchar i des and , i n s om e i n s t ances , p ro t e i n ANC s

T a b l e 1 ). H o w e v e r d u e t o t h e d i f f e re n t le g u m e s ,

ge rm i na t i on cond i t i ons and ana l y t i ca l m e t hods u s ed ,

t h e r e s u l t s o n t h e e f f e c t o f g e r m i n a t i o n o n l e g u m e

A N C s , c a n h a r d l y b e c o m p a r e d .

H e a t - t r e a t m e n t o f l e g u m e s e e d s n o t o n l y c a u s e s a

n u m b e r o f d e s i r a b le m o d i f ic a t i o n s t o t h e i r c h e m i c a l

com pos i t i on , nu t r i t i ona l and o rgano l ep t i c p roper t i e s

W u e t a l . , 1995 ) , bu t can a l s o i nduce l o s s es o f

es s en t i a l am i no ac i d s and v i t am i ns . Hea t dena t u ra -

t i o n a l s o h a s a m a r k e d i n f l u e n c e o n t h e

f u n c t i o n a l / t e c h n o l o g i c a l p r o p e r t i e s o f l e g u m e p r o -

t e i n s , wh i ch a re s t r i c t l y r e l a t ed t o t he i r m od i f i ed

phys i co -chem i ca l cha rac t e r i s t i c s , nam el y d i s s oc i a t i on

i n t o cons t i t uen t s ubun i t s , un fo l d i ng and s u r face ex -

p o s u r e o f t h e h y d r o p h o b i c s id e g r o u p s . H e a t d e n a t u -

r a t io n i s u s u a l l y a c c o m p a n i e d b y a d e c r e a s e o f s o l u-

b i l i t y , wh i ch r e s u l t s f rom t he agg rega t i on o f t he

u n f o l d e d m o l e c u l e s . T h e s e c h a n g e s a f f e c t a n u m b e r

o f func t i ona l p roper t i e s , s uch as t he ab i l i t y o f ge l i f i -

Table 1

Effect of soaking and cooking A) and germination B) on selected legum e seed ANCs. Values

amount/activity after treatment

re

given as percent residual

Genus TI Phytic acid Polyphenols

A B A B A B

Cajanus cajan 0 a 65 b 85 a 35 b 60 a 30 b

Canavalia ensiformis 0 60 c 70 c 35 65 c 20 c

Cicer arietinum

0 d 30 c 75 d 40 e 40 d 20 e

Glycine ma x 0 f 60 f 0 f 35 e 25 f 30 f

Lens culinaris variabilis 0 g 70 h 65 g 55 h 70 g 50 h

Lens culinaris vulgaris 0 g 75 h 70 g 30 h 60 g 50 h

Phaseolus mungo 0 i 40 i 80 i 40 i 35 i 25 i

Phaseolus vulgaris bla ck) 10 1 60 m 60 1 35 m 50 1 35 m

Phaseolus vulgaris

red ) 30 n 65 o 50 n 40 o 55 n 30 o

Pisum sativum 0 p 60 p 50 p 45 p 40 p 35 p

Psophocarpus tetragonobolus 0 q 70 r 50 q 40 r 80 q 30 r

Viciafaba 20 s 60 t 50 s 40 t 80 s 55 t

a Mulim ani and Para mjyot hi, 1993; b Wede r and Link, 1993; c Babar et al., 1988; d Attia et al., 1994; e Savag e and Thom pson , 1993; f Liu

and Markak is, 1987; g Vida l-V alv erde et al., 1994; h Batr a et al., 1986; i Redd y et al., 1978; i Des phan de and Chery an, 1984; m Trug o et

al., 1980; Dhurandh ar and Chang, 1990; o Nielsen and Liener, 1988; p Gatel, 1994; q de Lume n and Salamamat , 1980;

r

King and

Puwastien, 1987; s Aletor et al., 1994; t Rahma et al., 1987.


8/15

8


ca t i on (Mo i l , 1988 ) , rheo l og i ca l p rope r t i e s o f t he

ge l s , foam i ng and em u l s i f i ca t i on capac i t y , a s o f t en

s hown i n s oybean p ro t e i n s y s t em s (Yam ag i s h i e t a l . ,

1980 ; I awabuch i e t a l . , 1991a ; and I awabuch i e t a l . ,

1991b ) . Ge l fo rm a t i on i s an i m por t an t p rope r t y fo r

t h e u s e o f s o y b e a n p r o t e i n s i n f o o d s y s t e m s . D i f f e r -

e n t g e l s a r e f o r m e d w i t h t h e m a j o r s o y b e a n s t o r a g e

p ro t e i n s 11S and 7S g l obu l in s : t he ge ls fo rm ed by

t he 7S g l obu l i n a re t r an s pa ren t , con t r a ry t o t he tu rb i d

ge l s fo rm e d by t he l l S g l obu l i n ; t hes e l a tt e r ge l s

be i ng m uch ha rde r (Nakam ura e t a l . , 1986 ) . Th i s has

b e e n a t t r i b u t e d t o t h e f o r m a t i o n o f i n t e r m o l e c u l a r

d i s u l ph i de exch ange r ea c t i ons i n t he l l S g l obu l i n -

f o r m e d g e l s . I n t h e g e l f o r m a t i o n o f t h e m i x e d

s ys t em , t he 7S and 11S g l obu l i n s i n t e rac t non -cova-

l en t l y t o fo rm s o l ub l e agg rega t es (Nakam ura e t a l . ,

1986 ) . The ex t en t o f t h i s i n t e rac t i on i s a f f ec t ed by

t he p ropo r t i ons o f the t wo g l obu l i n s and t he i r s ubun i t

com pos i t i on , l ead i ng t o va r i ous ge l a t i on behav i ou r s .

S i n c e t h e c o m p o s i t i o n o f 1 1 S h e x a m e r s v a r i e s a m o n g

s oybean cu l t i va r s , i t i s pos s i b l e t o con t ro l t he r a t e o f

ge l a t i on and t he ge l p rope r t i e s o f s oybean p ro t e i n s

t h r o u g h s e l e c t i o n a n d / o r b r e e d i n g o f s o y b e a n c u l t i -

va r s by cons i de r i ng t he con t r i bu t i on o f each s ubun i t

t o t he f i na l p rope r t i e s o f t he ge l s (Nakam ura e t a l . ,

1985a and Nakam ura e t a l . , 1985b ; Za rkadas e t a l . ,

1994 ; Ko e t a l. , 1995 ) . The ex t r em e va r i ab i l i ty i n t he

cond i t i ons o f p ro t e i n dena t u ra t i on (pH, i on i c s t r eng t h ,

p r e s e n c e o f f r e e s u l p h y d r y l o r d i s u l p h y d e g r o u p s ,

t e m p e r a t u r e , h e a t i n g t i m e a n d r a t e o f c o o l i n g ) c a n

s i gn i f i can t l y a l te r t he func t i ona l i t y o f t he p ro t e i n and

c o n s e q u e n t l y t h e t e c h n o l o g i c a l p e r f o r m a n c e o f a d e -

r i v e d f o o d .

F rom t he p rac t i ca l v i ewpo i n t , t he cook i ng t i m e i s

an i m por t an t va r i ab l e t o cons i de r fo r t he u t i l i za t i on

o f a l egum e s eed , a s i t i s ev i dence d by t he w e l l

k n o w n h a r d - to - c o o k p ro b l e m i n c o m m o n b e a n s ( R e -

y e s - M o r e n o a n d P a r e d e s - L b p e z , 1 9 9 3) .

6 . Pos i t i ve e f fe c t s o f l e gu m e se e d s i n th e d i e t

L e g u m e s e e d c o m p o n e n t s m a y p l a y a p o s i t i v e

ro l e i n t he d i e t o f no rm al and even pa t ho l og i ca l

s ub j ec t s . F o r exam pl e , i t h as been s hown t ha t an

i n d i r e c t c o n t r o l o f p o l y a m i n e p r e s e n c e i n t h e g u t b y

l ec t i n s can s l ow down i n t e s t i na l t u rnou t g rowt h

(B ard ocz e t a l. , 1995 ) . O t he r s t ud i es have s how n t ha

i nc l u s i on o f beans i n t he d i e t have s om e p ro t ec t i ve

e f f e c t s in r a ts f e d a h y p e r c h o l e s t e r o l e m i c d i e t ( D e

Angel i s e t a l . , 1995) .

C e l i ac pa t i en t s , who s how a s eve re i n t o l e rance

res pons e t o food con t a i n i ng g l i ad i n and g l u t en i n

(Al l m ann e t a l. , 1992 ) , can s a fe l y be f ed wi t h beany

foods wh i ch con t a i n on l y a l bum i ns and g l obu l i n s .

N e w p r o c e s s i n g t e c h n o l o g i e s p r o d u c i n g a l t e r n a t i v e

f o o d s a r e b e i n g d e v i s e d . F o r e x a m p l e , p e a p r o t e i n s

c a n n o t f o r m t h e c o m p l e x s t r u c t u r e o f t h e w h e a

g l u t en (Adam s e t a l . , 1991 ; B ahnas s ey e t a l . , 1986 ) ,

bu t t he u s e o f em u l s i f i e r s , wh i ch f avou r t he i n t e rac -

t io n o f p r o t e in s w i th c a r b o h y d r a t e s , h a v e b e e n u s e d

t o i m p r o v e t h e s t r u c t u r e o f t h e d o u g h ( S c h u s t e r ,

1984 ; Kovacs and Gero , 1995 ) .

D r i e d l e g u m e s e e d s a r e k n o w n t o e l i c i t a l o w

g l ycem i c r e s pons e (Yoko t a e t a l . , 1994 ) , due t o

va r i ous f ac t o r s i nc l ud i ng a h i gh p ropo r t i on o f am y-

l o s e i n l egum e s t a rch , wh i ch i nc reas es t he p robab i l -

i t y o f t he i r r e t rog rada t i on , t he pos s i b l e i n t e rac t i ons o f

a m y l o l y t i c e n z y m e s w i t h a n t i n u t r i t i o n a l c o m p o u n d s

a n d t h e c o n t e n t a n d c o m p o s i t io n o f o l ig o s a c c h a r id e s

S l o w c a r b o h y d r a t e s c a n b e b e n e f i c i a l t o h e a l th a n d

he l p t o p reven t d i s eas es s uch as t ype I I d i abe t es ,

ca rd i ovas cu l a r p rob l em s and obes i t y . The l ow i n s u l i n

requ i r em en t i n t he pos t -p rand i a l phas e o f a m ea l r i ch

i n s l o w c a r b o h y d r a t e s c o u l d e x p l a in th e s e a p p a r e n

benef i c i a l e f f ec t s .

7 . D e v e l o p m e n t s a n d p e r s p e c t i v e s i n t h e p r o d u c

t i o n a n d u s e o f s e e d p r o t e i n s

S t udy o f s eed s t o rage p ro t e i n s and t he i r genes

have p l ayed an i m por t an t ro l e i n t he e s t ab l i s hm en t

a n d d e v e l o p m e n t o f p l a n t m o l e c u l a r b i o l o g y t e c h -

n i ques , m a i n l y due t o t he advan t ages p rov i de d by t he

m as s i ve s yn t hes i s o f f ew p ro t e i n s i n a s pec i f i c o rgan

d u r i n g s e e d d e v e l o p m e n t . A d e t a i l e d u n d e r s t a n d i n g

o f s t o rage p ro t e i n s t ruc t u re and d i ve r s i t y is an i m por -

t an t p re requ i s i t e fo r a t t em p t s t o m an i pu l a t e qua l i t y

becaus e i t i nd i ca t e s t he ex t en t t o wh i ch t he s t ruc t u re

o f t h e p r o t e i n c a n b e a l t e r e d w i t h o u t a f f e c t i n g s e e d

b i o l og y (S hew ry e t a l. , 1995 ) .

M a n y o f t h e s t o r a g e p r o t e i n g e n e s o f i m p o r t a n t

c r o p p l a n t s h a v e b e e n c l o n e d a n d c h a r a c t e r i z e d a n d


9/15

M. Duranti, C. Gius Field Crops Research 53 1997) 31-45 39

t he i r r egu l a t i on i nves t i ga t ed Go l dberg e t a l. , 1989 ;

S ho t we l l and Lark i n s , 1989 ; S h i r s a t , 1991 ) . Thes e

s t u d i e s h a v e p r o v i d e d a m o l e c u l a r b a s i s f o r t h e

i n t roduc t i on , apa r t f rom t he t r ad i t i ona l b reed i ng and

s e l ec t i on app roaches , o f p l an t gene t i c eng i nee r i ng .

G e n e r a l l y s p e a k in g , t h r e e g e n e t i c a p p r o a c h e s c a n b e

c o n s i d e r e d t o i m p r o v e t h e q u a l i t y o f l e g u m e p r o -

t e in s : i ) r egu l a t i on o f gene exp res s i on , i i ) i n t roduc-

t i on o f fo re i gn genes , and i i i) m od i f i ca t i on o f genes

enco d i ng a g i ven s t o rage p ro t e i n Tab e e t a l ., 1993 ) .

T h e p o s s i b i l i t y o f e x p r e s s i n g a n d a c c u m u l a t i n g

l egum e s t o rage p ro t e i n s i n t r an s gen i c p l an t s , s uch as

p e t u n i a Petunia hybrida), t o b a c c o Nicotiana

tabacum

L.) Be ac hy e t a l ., 1985; Br ay e t a l ., 1987;

Lawt on e t a l . , 1987 ) , Arabidopsis a n d Brassica

napus r ape s eed ) A l t enba ch e t a l ., 1989 ; C l e rcq e t

a l . , 1990a and C l e rcq e t a l . , 1990b ) have been ex -

p l o i t ed . Th i s app roach has o f t en m ade ava i l ab l e i n -

fo rm at i on on s eed p ro t e i n fo l d i ng , a s s em b l y , t r an s -

p o r t a n d d e p o s i t i o n , a n d i n m a n y c a s e s h a s a l l o w e d

t o i m p r o v e t h e c o n t e n t o f S - c o n t a i n i n g a m i n o a c id s ,

a s w i t h t he 2S s t o rage p ro t e i n s o f B raz i l nu t

Bertholletia excelsa)

1 8 m e t h i o n i n e r e s i d u e s / 1 0 0

am i no ac i d s ) A l t enb ach e t a l ., 1989 ; A l t enb ach e t

a l . , 1992; Saalbach e t a l . , 1995) . Now that the t rans-

f o r m a t i o n o f s o m e l e g u m e p l a n t s , n a m e l y Glycine

max a n d Vicia narbonensis, has been es t ab l i s hed

Saalbach e t a l . , 1994; Saalbach e t a l . , 1995) , th i s

a p p r o a c h w i l l b e c o m e o n e o f t h e m o s t p r o m i s i n g

m e t h o d s f o r i m p r o v i n g t h e n u t r i t i o n a l q u a l i t y o f

l egum e p ro t e i n s .

G e n e t r a n s f e r o f s e e d p r o t e i n s c o n f e r r i n g i n s e c t

r e s i s t ance i n t o c rop p l an t s i s an exc i t i ng goa l i n t he

f i e l d o f p l an t p ro t ec t i on . Exce l l en t cand i da t es fo r

t r ans fe r and exp res s i on a re s e r i ne p ro t e i nas e i n -

h ib i tors An et a l. , 1989; H i lder e t a l . , 1987) , l ec t ins

C hr i s pee l s and R a i khe l , 1991 ) and a -am y l as e i n -

h i b i t o r s Hu es i ng e t a l. , 1991 ; I s h i m o t o and Ki t a -

m ura , 1989 ; S ch roed er e t a l ., 1993 ) .

F r o m t h e v i e w p o i n t o f le g u m e p r o t e i n f u n c t io n a l -

i t y , a cha l l eng i ng p ro j ec t i s t he dom ai n -exchang i ng

gene t echno l ogy , cons i s t i ng i n t he cons t ruc t i on o f

n e w g e n e s o f a g i v e n p r o t e i n , b u t c o n t a in i n g s e l e c t e d

dom ai n s ) o f ano t he r s ubun i t . S i nce t he con t r i bu t i on

o f e a c h m o n o m e r i c u n i t o f a h e t e r o g e n e o u s s t o r a g e

p ro t e i n t o t he func t i ona l i t y depends upon t he p roper -

t i e s o f t h e m o n o m e r i t s e l f , t h e e x c h a n g e o f s o m e

c r i t i c a l d o m a i n w i l l a l l o w t h e p r o d u c t i o n o f n e w

s t o rage p ro t e i n s w i t h i m proved func t i ona l o r nu t r i -

t i ona l qua l i t i e s . A s oybean 11S g l obu l i n dom ai n -ex -

chang i ng gene can be eas i l y exp res s ed i n a t r an s -

g e n i c s o y b e a n p l a n t a s a s t a b l e m o n o m e r w h i c h c a n

s e l f - a s s em b l e to a 11S g l obu l i n m o l ecu l e U t s um i ,

1990) .

The va r i ab l e r eg i ons o f t he s t o rage p ro t e i n genes ,

wh i ch have l i t t l e func t i on i n fo rm i ng and m a i n t a i n -

i ng t he s t o rage p ro t e i n s t ruc t u re and m ay t o l e ra t e

m o d i f i ca t i on Ni e l s en , 1985 ) , a re the t a rge t s o f cho i ce

fo r t hes e t echn i ques . Thes e r eg i ons a re hyd roph i l i c

and a re t hus u s ua l l y l oca t ed on t he s u r f ace o f t he

p ro t e i n m o l ecu l e . There fo re , t he de l e t i on o f a po r t i on

o r a l l o f the va r i ab l e r eg i ons cau s es an i nc rea s e i n

h y d r o p h o b i c i t y o f t he m o n o m e r m o l e c u l e a s a w h o l e .

Th i s change a f f ec t s t he m o l ecu l a r s t ab i l i t y o f t he

p ro t e i n and , a s a r e s u l t , t he em u l s i fy i ng p rope r t y , the

ge l fo rm i ng r a t e , and t he ge l ha rdnes s a s s hown fo r

t he s oybean 11S p ro t e i n Ha que e t a l. , 1982 , Ha que

and Ki t o , 1982 ; Nakam ura e t a l . , 1984 ; Ka t o and

Yutan i , 1988; K im et a l. , 1990). The se m od ern

a p p r o a c h e s i m p l y a p r e f o r m e d k n o w l e d g e o f p r o t e i n

s t ruc t u re / s t ab i l i t y r e l a t i ons h i p s . When ava i l ab l e , 3D

s t ruc t u res o f s t o rage p ro t e i n s , un fo r t una t e l y an un -

d e r d e v e l o p e d f ie l d S u z u k i e t al ., 1 9 8 3; L a w r e n c e e t

a l . , 1990) , can be used as a t emplate to s imulate

m od i f i ca t i ons a i m ed a t i m prov i ng t he nu t r i t i ona l and

f u n c t io n a l p e r f o r m a n c e o f a s e e d p r o t e in s i m u l a t ed

m ut age nes i s ) Dy er e t a l ., 1993 ) .

8 C o n c l u s i o n s

D e s p i t e t h e i r a n c i e n t u s e f o r f o o d a n d f e e d a n d

t h e m a n y y e a r s o f i n te n s e r e s e a r c h, l e g u m e s e e d s a n d

t he i r com ponen t s s t i l l d es e rve fu r t he r exp l o i t a t i on .

The d i r ec t bene f i t s t o hum an nu t r i t i on and hea l t h ,

and t he po t en t i a l i t i e s o f t he p ro t e i n s and o t he r com -

pounds fo r func t i ona l i t y pu rpos es i n food a rch i t ec -

t u re , a s we l l a s t he i r r e l a t i ve i nexpens i venes s l a rge l y

c o m p e n s a t e t h e , o f t e n e a s i l y r e m o v a b l e , p r e s e n c e o f

an t i nu t r i t i ona l com pounds .

Af t e r t he i n t ens i ve u s e o f b i o t echno l og i ca l t ech -

n i ques i n t he f i e l d du r i ng t he l a s t decade , m any

s c i en ti s ts hav e r e cog n i zed t ha t a be t t e r u t i l i za ti on o f

l e g u m e s e e d s m a y a r i s e o n l y f r o m a h i g h l y i n t e -

g ra t ed app roach , whe re t he ro l e o f t he va r i ous s eed

com ponen t s bo t h i n t he s eed and i n ou r d i e t s has t o


10/15

4 0


be understood in detail and the desired modifications

developed accordingly

A c k n o w l e d g e m e n t s

Research supported by National Research Counci l

o f I ta ly Specia l Project RA ISA Subproject No. 2 .

R e f e r e n c e s

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o f b i o s y n t h e s i s , e x t r a c e ll u l a r s e c r e t i o n a n d i n t r ac e l l u la r s t o r a g e

o f p r o t e i n s i n p l a n t c e l l s . A n n u . R e v . P l a n t P h y s i o l . , 3 6 :

4 4 1 - 4 7 2 .

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Nu t r i t iona l and an t inu t r i t iona l cha rac te r i s t i c s o f se lec ted

Vicia

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s e e d p r o t e i n s b y t h e e x p r e s s i o n o f a c h i m e r i c g e n e e n c o d i n g a

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T h e i n f l u e n c e o f p e a s e e d t r y p s i n i n h i b i t o r s o n t h e i n v i t r o

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s e q u e n c e o f t h e 2 S s u l p h u r - r ic h p r o t e i n s f ro m s e e d s o f B r a z i l

n u t

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s i o n o f p r o t e i n a s e i n h i b i t o r s I a n d I I i n t r a n s g e n i c t o b a c c o

p l a n t s , e f f e c t s o n n a t u r a l d e f e n s e a g a i n s t

M. sexta

l a rvae .

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1 9 9 4 . E f f e c t o f c o o k i n g a n d d e c o r t i c a r i o n o n t h e p h y s i c a l

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c h i c k p e a

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