Planta
Planta 146, 575 577 (1979)
9 by Springer-Verlag 1979
Phytochrome and Internode Elongation in Chenopodiumpolyspermum L. The Light Fluence Rate during the Day and the End-of-day Effect A. L e c h a r n y and R. Jacques Laboratoire du Phytotron, C.N.R.S., F-91190 Gif-sur-Yvette, France
Abstract. The elongation of the f o u r t h internode o f fully green Chenopodiumpolyspermum L. is strongly stimulated by far-red light (FR) given at the end of the day. The end-of-day effect is m o r e i m p o r t a n t when the plants had been cultivated for several days with a main light period of 1 4 0 W m - 2 than with a main light period o f 85 W m 2. There exists a quantitative relationship between the F R end-of-day effect mediated by p h y t o c h r o m e and the value of the light fluence during the day.
site is not known. With the hypothesis that there is a regulation of the transport of trophic substances to the growing internode, the relationship between the value o f the energetic fluence o f the photosynthetically active radiations (PAR) and the end-of-day effect have been studied.
Key words: Chenopodium - D a y light fluence rate
The conditions of illumination to which the plants were submitted between their sowing and the FR treatment have been schematised in Fig. I. Tile temperature was 22~ C throughout the experiment. At the start of the FR treatment the fourth internode length of the plants was between 10 and 20 mm. The lengths attained at the end of the treatment were never those of fully expanded internodes. The organs were numbered according to their order of appearance after sowing, i.e., the third pair of leaves is the third pair above the cotyledons. The R and FR sources were described previously (Lecharny, 1979). An illumination of 6min at 60 Wm- 2 was saturating for the end-of-day reaction. It has been established that the effect observed can not be due to a calorific effect. The two values of PAR fluence rate have been chosen to avoid a significant difference of growth for 9 15 R between the batch given the higher PAR value (140 Wm-2, HWL treatment) and the batch given the lower value (85 Wm- 2, LWL treatment). The area of the leaves was measured with the aid of an automatic areameter (Hayashi denkoh Co. Ltd).
- E n d - o f - d a y effect - Internode elongation - Phytochrome.
Introduction The g r o w t h o f stems o f caulescent plants, cultivated in alternating light-darkness, depends on the quality of the light at the end of the day (Downs, 1957). The F R given just before darkness strongly stimulates the lengthening o f the internodes. In a preceding article (Lecharny, 1979) experiments leading to determination of the p h o t o r e c e p t i o n sites for F R in fully green Chenopodiurn polyspermum L. were discussed. T w o different types of organs were designated: the actual internode and the leaves just above and below the internode. The p h o t o t r a n s f o r m a t i o n of p h y t o c h r o m e is the primary reaction in the two organs. The photosynthetic organs are responsible for 60% o f the total effect observed when the entire plants are illuminated. The nature of the signal transmitted between the photoreceptive organs and the reaction
Abbreviations:D = darkness; FR = far-red light; HWL = white light at 140 Wm- 2; LWL = white light at 85 Win- z ; PAR = photosynthetically active radiation; R = red light; WL = white light
Material and Methods
Results and Conclusions The elongation o f the fourth internode of the plants in 9-15 D was the same for plants that had been given H W L treatment as for those given L W L treatment. In contrast, after a W L treatment o f 5 days there appeared a significant difference in the F R endof-day effect. The elongation stimulation o f the internodes by F R was stronger after a W L treatment at 140 W m - 2 than after a W L treatment at 85 W i n - 2 The difference increased with the length o f the W L treatment (Fig. 2). The light influences the growth
0032-0935/79/0146/0575/$01.00
576
A. Lecharny and R. Jacques: Phytochrome and Internode Elongation
1
9 W L d.15R " r - 9 w L d - 1 5 D -//'l. 9,vv Ld_ .~:Rel5D 4OWm 72 _
it
9wLa_l~R%,-I 85Wm-2b
L
II
dl "" t - - 9 W L d - 1 5 D 9 W L _ 5R - / / - - I d e 85Wm-2 I-. 9 W I ' _FR_15D W L treatment O; 5 ; 9 ; o r 13d
9
II
o/
f + FR
I
FR treatment 3d
2O (D
s
9
28
sowing
/
,6
3'1
W
days after sowing
+.~
rr" t_k. c-x...
-GI 10
Fig. 1. Experimental sequence of irradiations, a: ATLAS daylight fluorescent lamps and incandescent lamps; b: + 15 Wm -2 above 700 nm; c: PHILIPS TL 15 fluorescent lamps, 0.3 W m - 2; d : SYLVANIA Grolux WS fluorescent lamps; e: 6 min FR, 60 Win-2
E E 0.05). Highly significant differences (p < 0.01) were observed for the fresh weight, the dry weight, and for those values given per cm 2 of leaf. The fresh weight and the dry weight were highest for the stronger PAR fluence rate. The response to the increase in PAR fluence rate could be therefore due to an increase in the quantity of substrates produced by photosynthesis. The increase of PAR fluence rate did not stimulate the elongation of the internodes if PFR w a s present during the periods of darkness, i.e., the periods of greatest growth (Lecharny and Jacques, 1974). The phytochrome could therefore act by limiting to a certain value the availability of a growth substrate for the growing internodc. In the absence of PVR,the elongation of the internode could depend on the quantity of available substrate in the leaves.
577
References Dagnblie, P. : Th6orie et m6thodes statistiques. GembIoux: Duculot 1969
Downs, R.J. : Photoreversible control of elongation of Pinto beans and other plants under normal conditions of growth. Bot. Gaz. (Chicago) 118, 199-208 (1957) Grime, J.P., Jeffrey, D.W. : Seedling establishment in vertical gradients of sunlight. J. Ecol. 53, 621-642 (1965) Jacques, M., Jacques, R. : Floraison de deux plantes de jours longs : diversit~ de r6sponses aux 6clairements compl~mentaires rouge clair et rouge sombre, en fonction de la valeur de l'+clairement trophique. C.R. Acad. Sci. 287, 1333 1336 (I978) Lecharny, A. : Phytochrome and internode elongation in Chenopodium polyspermum L. Sites of photoreception. Planta 145, 405409 (1979) Lecharny, A., Jacques, R.: Phytochrome et croissance des tiges; variations de l'effet de la lumi6re en fonction du temps et du lieu de photoperception. Physiol. V+g. 12, 721 738 (1974) Morgan, D.C., Smith, H.: Linear relationship between phytochrome, photoequilibrium and growth in plants under simulated natural radiation. Nature 262, 210-212 (1976) Sanchez, R.A., Cogliatti, D. : The interaction between phytochrome and white light irradiance in the control of leaf shape in Taraxacure officinale. Bot. Gaz. (Chicago) 136, 281-285 (1975) Vince-Prue, D.: Photocontrol of stem elongation in light-grown plants of Fuchsia hybrida. Planta 133, 149 156 (1977) Received 21 March; accepted 5 June 1979
Planta 146, 575 577 (1979)
9 by Springer-Verlag 1979
Phytochrome and Internode Elongation in Chenopodiumpolyspermum L. The Light Fluence Rate during the Day and the End-of-day Effect A. L e c h a r n y and R. Jacques Laboratoire du Phytotron, C.N.R.S., F-91190 Gif-sur-Yvette, France
Abstract. The elongation of the f o u r t h internode o f fully green Chenopodiumpolyspermum L. is strongly stimulated by far-red light (FR) given at the end of the day. The end-of-day effect is m o r e i m p o r t a n t when the plants had been cultivated for several days with a main light period of 1 4 0 W m - 2 than with a main light period o f 85 W m 2. There exists a quantitative relationship between the F R end-of-day effect mediated by p h y t o c h r o m e and the value of the light fluence during the day.
site is not known. With the hypothesis that there is a regulation of the transport of trophic substances to the growing internode, the relationship between the value o f the energetic fluence o f the photosynthetically active radiations (PAR) and the end-of-day effect have been studied.
Key words: Chenopodium - D a y light fluence rate
The conditions of illumination to which the plants were submitted between their sowing and the FR treatment have been schematised in Fig. I. Tile temperature was 22~ C throughout the experiment. At the start of the FR treatment the fourth internode length of the plants was between 10 and 20 mm. The lengths attained at the end of the treatment were never those of fully expanded internodes. The organs were numbered according to their order of appearance after sowing, i.e., the third pair of leaves is the third pair above the cotyledons. The R and FR sources were described previously (Lecharny, 1979). An illumination of 6min at 60 Wm- 2 was saturating for the end-of-day reaction. It has been established that the effect observed can not be due to a calorific effect. The two values of PAR fluence rate have been chosen to avoid a significant difference of growth for 9 15 R between the batch given the higher PAR value (140 Wm-2, HWL treatment) and the batch given the lower value (85 Wm- 2, LWL treatment). The area of the leaves was measured with the aid of an automatic areameter (Hayashi denkoh Co. Ltd).
- E n d - o f - d a y effect - Internode elongation - Phytochrome.
Introduction The g r o w t h o f stems o f caulescent plants, cultivated in alternating light-darkness, depends on the quality of the light at the end of the day (Downs, 1957). The F R given just before darkness strongly stimulates the lengthening o f the internodes. In a preceding article (Lecharny, 1979) experiments leading to determination of the p h o t o r e c e p t i o n sites for F R in fully green Chenopodiurn polyspermum L. were discussed. T w o different types of organs were designated: the actual internode and the leaves just above and below the internode. The p h o t o t r a n s f o r m a t i o n of p h y t o c h r o m e is the primary reaction in the two organs. The photosynthetic organs are responsible for 60% o f the total effect observed when the entire plants are illuminated. The nature of the signal transmitted between the photoreceptive organs and the reaction
Abbreviations:D = darkness; FR = far-red light; HWL = white light at 140 Wm- 2; LWL = white light at 85 Win- z ; PAR = photosynthetically active radiation; R = red light; WL = white light
Material and Methods
Results and Conclusions The elongation o f the fourth internode of the plants in 9-15 D was the same for plants that had been given H W L treatment as for those given L W L treatment. In contrast, after a W L treatment o f 5 days there appeared a significant difference in the F R endof-day effect. The elongation stimulation o f the internodes by F R was stronger after a W L treatment at 140 W m - 2 than after a W L treatment at 85 W i n - 2 The difference increased with the length o f the W L treatment (Fig. 2). The light influences the growth
0032-0935/79/0146/0575/$01.00
576
A. Lecharny and R. Jacques: Phytochrome and Internode Elongation
1
9 W L d.15R " r - 9 w L d - 1 5 D -//'l. 9,vv Ld_ .~:Rel5D 4OWm 72 _
it
9wLa_l~R%,-I 85Wm-2b
L
II
dl "" t - - 9 W L d - 1 5 D 9 W L _ 5R - / / - - I d e 85Wm-2 I-. 9 W I ' _FR_15D W L treatment O; 5 ; 9 ; o r 13d
9
II
o/
f + FR
I
FR treatment 3d
2O (D
s
9
28
sowing
/
,6
3'1
W
days after sowing
+.~
rr" t_k. c-x...
-GI 10
Fig. 1. Experimental sequence of irradiations, a: ATLAS daylight fluorescent lamps and incandescent lamps; b: + 15 Wm -2 above 700 nm; c: PHILIPS TL 15 fluorescent lamps, 0.3 W m - 2; d : SYLVANIA Grolux WS fluorescent lamps; e: 6 min FR, 60 Win-2
E E 0.05). Highly significant differences (p < 0.01) were observed for the fresh weight, the dry weight, and for those values given per cm 2 of leaf. The fresh weight and the dry weight were highest for the stronger PAR fluence rate. The response to the increase in PAR fluence rate could be therefore due to an increase in the quantity of substrates produced by photosynthesis. The increase of PAR fluence rate did not stimulate the elongation of the internodes if PFR w a s present during the periods of darkness, i.e., the periods of greatest growth (Lecharny and Jacques, 1974). The phytochrome could therefore act by limiting to a certain value the availability of a growth substrate for the growing internodc. In the absence of PVR,the elongation of the internode could depend on the quantity of available substrate in the leaves.
577
References Dagnblie, P. : Th6orie et m6thodes statistiques. GembIoux: Duculot 1969
Downs, R.J. : Photoreversible control of elongation of Pinto beans and other plants under normal conditions of growth. Bot. Gaz. (Chicago) 118, 199-208 (1957) Grime, J.P., Jeffrey, D.W. : Seedling establishment in vertical gradients of sunlight. J. Ecol. 53, 621-642 (1965) Jacques, M., Jacques, R. : Floraison de deux plantes de jours longs : diversit~ de r6sponses aux 6clairements compl~mentaires rouge clair et rouge sombre, en fonction de la valeur de l'+clairement trophique. C.R. Acad. Sci. 287, 1333 1336 (I978) Lecharny, A. : Phytochrome and internode elongation in Chenopodium polyspermum L. Sites of photoreception. Planta 145, 405409 (1979) Lecharny, A., Jacques, R.: Phytochrome et croissance des tiges; variations de l'effet de la lumi6re en fonction du temps et du lieu de photoperception. Physiol. V+g. 12, 721 738 (1974) Morgan, D.C., Smith, H.: Linear relationship between phytochrome, photoequilibrium and growth in plants under simulated natural radiation. Nature 262, 210-212 (1976) Sanchez, R.A., Cogliatti, D. : The interaction between phytochrome and white light irradiance in the control of leaf shape in Taraxacure officinale. Bot. Gaz. (Chicago) 136, 281-285 (1975) Vince-Prue, D.: Photocontrol of stem elongation in light-grown plants of Fuchsia hybrida. Planta 133, 149 156 (1977) Received 21 March; accepted 5 June 1979
