Post Harvest Responses and Plant Growth Regulators

vice versa. Finally, I briefly .... plant tissues and organs are developmental stages in the life cycle and, like ... centrations at a given stage of ...
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15 Post Harvest Responses and Plant Growth Regulators MORRIS L I E B E R M A N

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Post Harvest Plant Physiology Laboratory, Beltsville Agricultural Research Center, Agricultural Research Service, U.S.D.A., Beltsville, Md. 20705

Regulation and control of post harvest metabolism with respect to ripening, aging, and senescence, has for many years been associated with degradation and the action of ethylene (1). Recent concepts and interpretations however, suggest that aging and senescence i n plant tissues are not only deteriorative processes but also developmental processes i n which other growth regulators play important roles (2). Thus, although ethylene is still considered a major influence on post harvest metabolism, the other plant hormones, the auxins, gibberellins, cytokinins, and abscisic acid, are also thought to s i g n i f i c a n t l y influence the aging process. Most likely, ethylene action results from interactions with these hormones. In this report I b r i e f l y review c l a s s i c a l observations on the effect of ethylene on post harvest tissues, especially those of f r u i t , and point out why ethylene was considered the ripening hormone. Secondly, I call attention to reasons for questioning ethylene as the exclusive ripening hormone and review recent data linking ethylene action to the action of other hormones and vice versa. F i n a l l y , I b r i e f l y discuss ethylene production and i n h i b i t i o n in plant tissue. These considerations may give r i s e to new concepts of controlling aging and senescence of plant tissues and of preserving crops after harvest. Physiological Responses to Ethylene. C l a s s i c a l l y , two types of f r u i t have been recognized with respect to their response to ethylene (I): (a) climacteric f r u i t , such as apples or avocado, which show an immediate r i s e i n respiration and an accelerated ripening rate when exposed to a few parts per m i l l i o n of ethylene; and (b) non-climacteric fruit, such as c i t r u s , which show a r i s e i n respiration during exposure to much higher concentrations of ethylene (100 ppm or more) then a return to the normal rate when ethylene is removed (Figure 1). Continuous application of such high levels of ethylene nevertheless accelerates ripening in non-climacteric fruit, which appear to r e s i s t reaction to ethylene (3). In contrast climacteric f r u i t 280

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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Air Air-} ethylene _I

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Days

0

2 4

I 6

I 8

L _ . - L _1_ ..L _L__J 10 12 14 16 18 20

Weeks Endeavor

Figure 1. Effect of ethylene on respiration of climacteric and nonclimacteric fruit. Ethylene causes greatest response in climacteric fruit when applied to mature fruit prior to the climacteric rise. In nonclimacteric fruit high concentrations of ethylene stimuhte respiration for short time periods. This stimulation is observed at any time upon application of ethylene (3).

u s u a l l y r e q u i r e o n l y a f e w ppm o f e t h y l e n e t o t r i g g e r t h e r i p ­ ening process. E t h y l e n e was b e l i e v e d t o be t h e r i p e n i n g hormone, b e c a u s e i t i s a n a t u r a l m e t a b o l i t e , i s v i r t u a l l y absent i n mature b u t n o n - r i p e n i n g f r u i t , and i s p r o d u c e d i n i n c r e a s i n g amounts j u s t p r i o r t o the o n s e t o f r i p e n i n g . Furthermore, ethylene applied a t r e l a t i v e l y low c o n c e n t r a t i o n s c a n i n d u c e r i p e n i n g a n d a g i n g i n green mature f r u i t , a c c e l e r a t i n g , but not a l t e r i n g , t h e n a t u r a l p r o c e s s e s . Compared t o r e l a t e d compounds, e t h y l e n e i s unique i n i t s e f f e c t i v e n e s s i n i n d u c i n g r i p e n i n g and a g i n g (Table I ) .

T a b l e I . Comparative E f f e c t i v e n e s s o f E t h y l e n e and R e l a t e d A n a l o g u e s i n P e a S t e m - S e c t i o n A s s a y (From B u r g and Burg (4) Compound Ethylene Propylene Vinyl chloride Carbon Monoxide Acetylene 1-Butene

Relative Activity;Moles/unit

effectiveness

1 130 2,370 2,900 12,500 140,000

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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The c l i m a c t e r i c r i s e i n r e s p i r a t i o n i s c o n s i d e r e d a p o r t e n d of the s h i f t i n metabolism from anabolism to c a t a b o l i s m . This t r a n s i t i o n from the f u l l y mature s t a t e t o the r i p e n i n g s t a t e o c c u r s more s h a r p l y i n f a s t e r g r o w i n g f r u i t , s u c h as t h e a p p l e , i n w h i c h t h e r e i s h y d r o l y t i c c o n v e r s i o n o f s t a r c h t o s u g a r s and of i n s o l u b l e p e c t i n s t o s o l u b l e p e c t i n s . There i s a l s o a l o s s o f c h l o r o p h y l l , and a s y n t h e s i s o f a n t h o c y a n i n s , c a r o t e n o i d s and x a n t h o p h y l l s , as w e l l as o c c u r r e n c e o f o t h e r s u c h r e a c t i o n s a s s o c i a t e d w i t h r i p e n i n g . I n n o n - c l i m a c t e r i c f r u i t , s u c h as c i t r u s , w h i c h show no c o m p a r a b l e r i s e i n r e s p i r a t i o n o r e t h y l e n e p r o d u c t i o n , g r o w t h and d e v e l o p m e n t a r e p r o l o n g e d and r i p e n i n g o c c u r s o n l y on t h e t r e e . An o r a n g e r e q u i r e s 8-11 months f r o m f u l l b l o o m t o m a t u r i t y , i n c o n t r a s t t o an a p p l e , w h i c h may r e q u i r e o n l y 4-5 months (5^) . R e s i s t a n c e t o E t h y l e n e as a R i p e n i n g A g e n t . The c l a s s i c a l concept of r i p e n i n g , e s p e c i a l l y of c l i m a c t e r i c f r u i t , i s t h a t e t h y l e n e t r i g g e r s a c a s c a d e o f r e a c t i o n s l e a d i n g t o r i p e n i n g and aging. T h i s c o n c e p t has b e e n q u e s t i o n e d b e c a u s e s u c h a t r i g ­ g e r i n g phenomenon i s a b s e n t i n n o n - c l i m a c t e r i c f r u i t . Also, u n d e r some c o n d i t i o n s e t h y l e n e does n o t t r i g g e r f r u i t r i p e n i n g , e v e n i n c l i m a c t e r i c f r u i t (6). T h u s , f o r e x a m p l e , a v o c a d o s do n o t r i p e n on t h e t r e e , d e s p i t e t h e i r i n t e r n a l a t m o s p h e r e o f a b o u t 0.1 ppm e t h y l e n e , a c o n c e n t r a t i o n w h i c h c a n i n d u c e r i p e n i n g i n the h a r v e s t e d f r u i t . Even t r e a t m e n t of the u n h a r v e s t e d fruit w i t h 50 ppm e t h y l e n e f o r 48 h o u r s does n o t c a u s e r i p e n i n g . The r e s i s t a n c e t o r i p e n i n g by e t h y l e n e e x t e n d s t o n e w l y h a r v e s t e d avocado f r u i t . A p p l i c a t i o n o f 100 ppm e t h y l e n e 1 h o u r a f t e r h a r v e s t had no e f f e c t on r i p e n i n g (7^). However 24 h o u r s a f t e r h a r v e s t , ethylene treatment c o n s i d e r a b l y a c c e l e r a t e d r i p e n i n g , much as e x p e c t e d . Such d a t a gave r i s e t o t h e i d e a o f an " a n t i r i p e n i n g " i n h i b i t o r , w h i c h , p r e s u m a b l y , i s most a c t i v e i n t h e u n h a r v e s t e d f r u i t and d i s s i p a t e d a f t e r h a r v e s t . G r a p e s a l s o do n o t r e s p o n d t o e t h y l e n e as e x p e c t e d . In d e v e l o p m e n t o f t h e g r a p e 3 s t a g e s o f g r o w t h c a n be d i s t i n g u i s h e d . An e a r l y r a p i d e n l a r g e m e n t i s f o l l o w e d by a s l o w s t a g e o f g r o w t h w h i c h i s a g a i n f o l l o w e d by a r a p i d g r o w t h s t a g e . The grape r i p e n s d u r i n g the t h i r d stage but w i t h o u t i n c r e a s e i n e t h y l e n e p r o d u c t i o n ( F i g u r e 2 ) . However, a s h a r p i n c r e a s e i n a b s c i s i c a c i d (ABA) does o c c u r and i s c o r r e l a t e d w i t h r i p e n i n g of the b e r r y . While the s e n s i t i v i t y t o e t h y l e n e i n c r e a s e d d u r i n g r i p e n i n g o f t h i s f r u i t (9) i t i s a l s o p o s s i b l e t h a t e t h y l e n e i s not the major r i p e n i n g f a c t o r . Such examples o f " a n o m a l o u s " r i p e n i n g b e h a v i o r d a t a s u g g e s t that other f a c t o r s i n a d d i t i o n to ethylene s i g n i f i c a n t l y a f f e c t r i p e n i n g and s e n e s c e n c e . The e x p e r i m e n t s w i t h c i t r u s and a v o c a d o s u g g e s t a n t i - r i p e n i n g s u b s t a n c e s and t h e s t u d i e s w i t h t h e d e v e l o p i n g g r a p e s u g g e s t t h a t ABA may p l a y a r o l e i n t h e s e processes. However, ABA a p p e a r s t o be a s u b s t i t u t e o r s u p p l e m e n t t o e t h y l e n e and n o t a t r i g g e r i n g a g e n t ( 1 0 ) .

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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VERAISON Plant Physiology

Figure 2. Ethylene concentration in extracted gas, weight of abscisic acid (ABA), fresh weight and volume per berry in "Doradillo" grapes during the stationary and ripening stages of growth (pre and post veraison) (S)

Other F a c t o r s I n f l u e n c i n g R i p e n i n g and Aging. Tradition­ a l l y , p l a n t hormones a r e a s s o c i a t e d w i t h g r o w t h and d e v e l o p m e n t of young v i g o r o u s t i s s u e s . I t i s now e v i d e n t , h o w e v e r , t h a t t h e s e hormones ( a u x i n s , c y t o k i n i n s , g i b b e r e l l i n s a n d ABA) may a l s o be i m p o r t a n t t o r i p e n i n g , a g i n g a n d s e n e s c e n c e , a n d t o many other aspects o f post harvest metabolism. Conversely, ethylene, w h i c h was a s s o c i a t e d w i t h r i p e n i n g i n a g i n g c e l l s a n d t i s s u e s , appears a l s o t o p l a y an i m p o r t a n t r o l e i n young v i g o r o u s l y growing t i s s u e s (11). W i t h o u t i n any way d i m i n i s h i n g t h e i m p o r t a n c e o f e t h y l e n e i n the c o n t r o l and r e g u l a t i o n o f r i p e n i n g and a g i n g , I emphasize the c r i t i c a l s u p p l e m e n t a l importance o f a u x i n s , g i b b e r e l l i n s , and

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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c y t o k i n i n s , a l o n g w i t h ABA, i n t h e r i p e n i n g and a g i n g p r o c e s s e s i n post h a r v e s t t i s s u e s . R i p e n i n g , a g i n g , and s e n e s c e n c e o f p l a n t t i s s u e s and o r g a n s a r e d e v e l o p m e n t a l s t a g e s i n t h e l i f e c y c l e and, l i k e any o t h e r s t a g e i n g r o w t h and d e v e l o p m e n t , a r e r e g u l a t e d and c o n t r o l l e d , a t t h e o r g a n i z a t i o n a l l e v e l , by i n t e r ­ a c t i o n between e t h y l e n e and o t h e r p l a n t hormones. Evidence f o r Hormonal I n t e r a c t i o n s w i t h E t h y l e n e . There i s e v i d e n c e o f a n t a g o n i s m b e t w e e n a u x i n s , g i b b e r e l l i n s and c y t o k i n i n s on one hand and e t h y l e n e and ABA on t h e o t h e r ( 1 2 ) . The c o n t r o l o f f r u i t g r o w t h , d e v e l o p m e n t , r i p e n i n g and a g i n g may depend on t h e r e l a t i v e i m p o r t a n c e o f a s p e c i f i c hormone i n the t o t a l hormonal balance. V a r i o u s hormones may t e n d t o be d o m i n a n t o r l a t e n t d e p e n d i n g p r o b a b l y on t h e i r l e v e l s o r c o n ­ c e n t r a t i o n s a t a g i v e n stage of the l i f e c y c l e . F i g u r e 3 shows t h e h y p o t h e t i c a l k i n e t i c s o f g r o w t h , r e s p i r a t i o n and r e l a t i v e hormone l e v e l s i n a c l i m a c t e r i c f r u i t at d i f f e r e n t stages of i t s l i f e c y c l e . H y p o t h e t i c a l hormone l e v e l s d u r i n g d e v e l o p m e n t and r i p e n i n g h a v e b e e n s p e c u l a t e d on b e f o r e ( 1 3 ) . The r a t i o n a l e f o r t h i s o u t l i n e i s b a s e d on t h e known i n f l u e n c e s o f t h e v a r i o u s hormones on c e l l d i v i s i o n ,

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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e l o n g a t i o n , and senescence. I n v e r y young f r u i t , c e l l d i v i s i o n i s t h e m a j o r a c t i v i t y , and, a u x i n s , g i b b e r e l l i n s a n d c y t o k i n i n s a r e a t t h e i r h i g h e s t l e v e l s , e t h y l e n e i s v i r t u a l l y a b s e n t , and ABA c o n t e n t i s r e l a t i v e l y h i g h ( 1 4 ) . The l e v e l s o f hormones r e f l e c t t h e i r a c t i v i t i e s which are presumably h i g h d u r i n g c e l l division. ABA may o p e r a t e as a b r a k e d u r i n g t h i s s t a g e b y opposing the p o s s i b l e e x c e s s i v e growth e f f e c t s o f h i g h con­ c e n t r a t i o n s o f some o f t h e s e hormones. A l s o , ABA i s somehow a l s o r e l a t e d t o w a t e r uptake w h i c h i s i m p o r t a n t i n young t i s s u e s (15). D u r i n g r a p i d c e l l e l o n g a t i o n g i b b e r e l l i n s may t e n d t o i n c r e a s e somewhat i n k e e p i n g w i t h t h e i r i m p o r t a n c e i n e l o n g a t i o n processes. During m a t u r a t i o n , t h e a u x i n s , g i b b e r e l l i n s , and c y t o k i n i n s d e c l i n e , r e a c h i n g v e r y low l e v e l s toward the end o f the m a t u r a t i o n p e r i o d . I t i s d u r i n g t h i s time t h a t t h e l e v e l s o f e t h y l e n e a n d ABA b e g i n t o r i s e , p r e c e d i n g somewhat t h e increase i n r e s p i r a t i o n associated with climacteric f r u i t . A l t h o u g h o n l y l i m i t e d d a t a a r e a v a i l a b l e o n endogenous hormone l e v e l s d u r i n g d i f f e r e n t s t a g e s o f f r u i t g r o w t h a n d d e v e l o p m e n t , some d a t a do s u p p o r t t h i s s i m p l i s t i c m o d e l . F o r example stem g r o w t h a p p e a r s t o r e s u l t f r o m a r i s e i n a u x i n l e v e l s (16) and r a t e o f tomato f r u i t r i p e n i n g i s i n v e r s e l y r e ­ l a t e d t o c y t o k i n i n content (17). Evidence f o r the i n t e r ­ r e l a t i o n s h i p s o f h o r m o n a l l e v e l s a n d r i p e n i n g and a g i n g i s a l s o o b t a i n e d f r o m e x p e r i m e n t s i n w h i c h v a r i o u s hormones a r e added t o r i p e n i n g o r aging f r u i t t i s s u e s . For example, t h e a u x i n , β-naphthylacetic a c i d , h a s l i t t l e e f f e c t o n c o l o r d e v e l o p m e n t i n r i p e n i n g banana p e e l d i s k s , w h e r e a s t h e c y t o k i n i n b e n z y l a d e n i n e c o n s i d e r a b l y r e t a r d s c o l o r i n g and, t h e r e f o r e , r i p e n i n g i n t h e s e d i s k s ( 1 8 ) . The t e n d e n c y o f exogenous a u x i n t o r e t a r d r i p e n i n g may be c o u n t e r b a l a n c e d b y t h e a b i l i t y o f a u x i n s t o s t i m u l a t e ethylene production. G i b b e r e l l i n s also retard r i p e n i n g (color f o r m a t i o n ) i n banana p e e l d i s k s , w h e r e a s a p p l i e d ABA a t 10"5 t o 10"^M a c c e l e r a t e s r i p e n i n g , as m i g h t be e x p e c t e d f r o m t h e known a n t a g o n i s m b e t w e e n ABA and g i b b e r e l l i n s ( 1 8 ) . The i n f l u e n c e o f g r o w t h hormones ( a u x i n s , c y t o k i n i n s , a n d g i b b e r e l l i n s ) a n d ABA o n e t h y l e n e p r o d u c t i o n i n a p p l e t i s s u e s l i c e s o f v a r i o u s s t a g e s o f m a t u r i t y b e f o r e , d u r i n g , and a f t e r t h e c l i m a c t e r i c r i s e i n r e s p i r a t i o n i s shown i n F i g u r e 4. P r e c l i m a c t e r i c t i s s u e s l i c e s , w h i c h e v o l v e v i r t u a l l y no e t h y l e n e , are s t r o n g l y i n h i b i t e d by the c y t o k i n i n , i s o p e n t e n y l adenosine ( I P A ) , i n d o l e a c e t i c a c i d ( I A A ) , and t o a l e s s e r e x t e n t b y g i b b e r e l l i c a c i d (GA). The e f f e c t o f a l l t h r e e hormones i s e v e n more i n h i b i t i n g t o e t h y l e n e p r o d u c t i o n b y t h e s e t i s s u e s . However, ABA s t i m u l a t e s e t h y l e n e p r o d u c t i o n i n p r e c l i m a c t e r i c t i s s u e slices. A t l a t e r s t a g e s i n t h e r i p e n i n g p r o c e s s , IAA and GA do n o t i n h i b i t e t h y l e n e p r o d u c t i o n a n d IAA may a c t u a l l y s t i m u l a t e . On t h e o t i i e r h a n d , IPA c o n s i s t e n t l y i n h i b i t s e t h y l e n e p r o d u c t i o n a t a l l s t a g e s o f r i p e n i n g t h r o u g h o u t t h e c l i m a c t e r i c and p o s t ­ c l i m a c t e r i c p e r i o d s . No g r e a t e r r e t a r d a t i o n i s a c h i e v e d b y a d d i t i o n o f GA a n d IAA t o IPA a t t h i s s t a g e .

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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Preclimactenc

CENTURY

Early climacteric rise

Figure 4. Influence of IAA, GA, cytokinin (IPA), and ABA, alone and in combinations, on ethylene production of pre-climateric, climacteric, and post-climateric apple tissue slices

•—control • —IAA • —GA A—IPA

O—ABA

Π — I A A + IPA Δ — I A A + IPA + G A

ABA s t i m u l a t e s e t h y l e n e p r o d u c t i o n i n p r e c l i m a c t e r i c and e a r l y c l i m a c t e r i c t i s s u e s l i c e s b u t s u b s e q u e n t l y has l i t t l e e f f e c t on e t h y l e n e p r o d u c t i o n i n a g i n g t i s s u e s l i c e s f r o m a p p l e s . These d a t a a l s o show t h a t t h e i n f l u e n c e o f a hormone may be c o n s i d e r a b l y a l t e r e d b y c o m b i n a t i o n s w i t h o t h e r hormones. Thus, IAA, w h i c h s t i m u l a t e s e t h y l e n e p r o d u c t i o n i n c l i m a c t e r i c and p o s t c l i m a c t e r i c t i s s u e does n o t s t i m u l a t e b u t i n h i b i t s when c o m b i n e d w i t h IPA a n d GA. F e e d b a c k R e l a t i o n s h i p B e t w e e n E t h y l e n e and O t h e r P l a n t Hormones. I f e t h y l e n e p r o d u c t i o n i n r i p e n i n g f r u i t i s an i n d e x o f a g i n g and s e n e s c e n c e , t h e n i t s s u p p r e s s i o n s h o u l d r e s u l t i n r e t a r d a t i o n , o r antagonism t o r i p e n i n g , a g i n g , and senescence.

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

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15.

LiEBERMAN

Post Harvest

Responses

287

At t h e p r e c l i m a c t e r i c stage o f development, j u s t p r i o r t o t h e r i s e i n ethylene production,the concentrations of auxins, g i b b e r e l l i n s , a n d c y t o k i n i n s a r e assumed t o be v e r y l o w . A d d i t i o n o f l a r g e amounts o f t h e s e g r o w t h hormones t e n d t o s u p p r e s s e t h y l e n e p r o d u c t i o n . However, a f t e r e t h y l e n e p r o d u c t i o n s t a r t s and a c c e l e r a t e s , o n l y c y t o k i n i n c o n s i s t e n t l y s u p p r e s s e s ethylene p r o d u c t i o n . This suggests a s p e c i a l antagonism be­ tween c y t o k i n i n s and e t h y l e n e p r o d u c t i o n . T h i s a n t a g o n i s m i s c o n s i s t e n t w i t h t h e w e l l known r e t a r d a t i o n e f f e c t o f c y t o k i n i n s on l o s s o f c h l o r o p h y l l and p r o t e i n i n a g i n g l e a v e s ( 1 9 ) . The r e l a t i v e l a r g e s c a l e p r o d u c t i o n o f e t h y l e n e b y a g i n g t i s s u e s , a f t e r the r i p e n i n g r e a c t i o n s a r e f u l l y i n motion, r a i s e s t h e q u e s t i o n as t o w h e t h e r o r n o t c o n t i n u o u s p r e s e n c e o f e t h y l e n e i s n e c e s s a r y f o r t h e p r o g r e s s o f a g i n g and s e n e s c e n c e . F a i r l y l a r g e amounts o f e t h y l e n e a r e p r o d u c e d e v e n f r o m f u l l y senescent t i s s u e s . This might suggest a l o s s o f c o n t r o l over e t h y l e n e p r o d u c t i o n . Such a n i n t e r p r e t a t i o n w o u l d mean t h a t e t h y ­ lene produced by post c l i m a c t e r i c t i s s u e i s a by-product o f ag­ i n g metabolism wherein c o n t r o l o f hormonal s y n t h e s i s i s l o s t . However, t h e f a c t t h a t e t h y l e n e p r o d u c t i o n i n p o s t - c l i m a c t e r i c t i s s u e s c a n be s u p p r e s s e d b y a c y t o k i n i n s u g g e s t s t h a t i t i s a l w a y s u n d e r p h y s i o l o g i c a l c o n t r o l and h o r m o n a l l y r e g u l a t e d . Scheme f o r I n t e r r e l a t i o n s h i p b e t w e e n E t h y l e n e a n d O t h e r Hormones. From t h e s e d a t a and o t h e r s p r e s e n t e d b e l o w , one c a n a r r i v e a t a s i m p l e scheme f o r t h e a n t a g o n i s t i c and s u p p o r t i v e r e l a t i o n s h i p s o f t h e s e hormones i n m e t a b o l i s m . F i g u r e 5 shows h y p o t h e t i c a l i n t e r c o n n e c t i o n s between t h e v a r i o u s hormones as t h e y r e l a t e t o c e l l d i v i s i o n , g r o w t h , d e v e l o p m e n t and senescence d u r i n g the l i f e c y c l e o f a p l a n t o r organ. Two c a t e g o r i e s o f hormones may be d i s t i n g u i s h e d : ( 1 ) t h e a u x i n s ^ g i b b e r e l l i n s , and c y t o k i n i n s , w h i c h a r e m a i n l y a s s o c i a t e d w i t h growth and development by r e g u l a t i n g c e l l d i v i s i o n , e n l a r g e ­ ment and m a t u r a t i o n , and (2) e t h y l e n e a n d ABA, w h i c h g e n e r a l l y t e n d t o oppose o r a n t a g o n i z e t h e a c t i v i t i e s o f c a t e g o r y 1 h o r ­ mones and t o f u n c t i o n m a i n l y i n s e n e s c e n c e and a g i n g . During g r o w t h and d e v e l o p m e n t c a t e g o r y 2 hormones may oppose t h e e x c e s s i v e a c t i o n s o f c a t e g o r y 1 hormones, w h i c h may o t h e r w i s e c a u s e d i s t o r t e d and a b n o r m a l g r o w t h e f f e c t s . These hormones may e x i s t i n f e e d b a c k l o o p s . A u x i n s a r e known t o s t i m u l a t e e t h y l e n e p r o d u c t i o n (20) and, c o n v e r s e l y , e t h y l e n e i s known t o r e d u c e a u x i n l e v e l s (21) ( 2 2 ) . G i b b e r e l l i n s and ABA a r e known t o oppose e a c h o t h e r i n t h e i r i n f l u e n c e on i n d u c t i o n o f α-amylase s y n t h e s i s i n t h e b a r l e y a l e u r o n e l a y e r ( 2 3 ) . C y t o k i n i n s and ABA a r e known t o oppose e a c h o t h e r i n t r a n s p i r a t i o n phenomena w i t h r e s p e c t t o r e g u l a t i n g o p e n i n g and c l o s i n g o f s t o m a t a ( 2 4 ) . C y t o k i n i n i s known t o b o t h s t i m u l a t e (25) and d e p r e s s ( 2 6 ) e t h y l e n e p r o d u c t i o n i n p l a n t s . I do n o t know o f a n o p p o s i n g a c t i o n o f e t h y l e n e on c y t o k i n i n s y n t h e s i s o r a c t i v i t y b u t s u c h may v e r y l i k e l y e x i s t .

Plimmer et al.; Pesticide Chemistry in the 20th Century ACS Symposium Series; American Chemical Society: Washington, DC, 1977.

288

PESTICIDE C H E M I S T R Y IN T H E 2 0 T H

CENTURY

INTERACTIONS BETWEEN PLANT HORMONES

FEEDBACK

LOOPS

CATEGORY 2

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CATEGORY I

->CELL

ETHYLENE

DIVISION