Scientia Itorticulturae, 6 (1977) 15--26 15

Elsevier Scientific Publishing Company, Amsterdam -- Printed in The Netherlands

EFFECT OF LIGHT CONDITIONS ON THE DEVELOPMENT OF THE

INFLORESCENCE IN TOMATO

J.M. KINET
Centre de Physiologie V$g$tale Appliqu$e (I.R.S.I.A.), D~partement de Botanique, Uni-
versit~ de Liege, Sart Tilman, B-4000 LiSge (Belgium)
(Received 15 June 1976)

ABSTRACT

Kinet, J.M., 1977. Effect of light conditions on the development of the inflorescence in
tomato. Scientia Hort., 6: 15--26.

By growing tomato plants (Lycopersicon esculentum Mill. ) in 4 different light regimes

(2 photoperiods -- 8 and 16 h -- combined with 2 light intensities -- 9,000 and 18,000 ergs
cm -2 s-' ), it was shown that increasing light integrals hastened flower initiation, greatly
promoted the development of the inflorescence and increased the rates o f leaf production
and the growth of the stem. In similar light integrals, flower initiation was earlier and in-
florescence development far better in short photoperiods than in long ones; the rates of leaf
production were almost the same and stem growth was greater in long days.
Transfer experiments from favourable to insufficient light conditions and inverse
transfers at different times during the life of the plant indicated that light conditions were
critical at the time of, and after, the macroscopic appearance of the inflorescence. At that
stage, a transfer to low light conditions for 10 days induced complete abortion of the truss
in our growth conditions. The effect of a transfer from insufficient to favourable light
conditions was slower since at least 15 days in these latter conditions were required in
order to achieve the development of the inflorescence.

INTRODUCTION

Several studies have stressed the difficulty in obtaining satisfactory develop-

ment of the first inflorescence in early tomato growing in temperate countries
such as the United Kingdom (Cooper, 1964; Cooper and Hurd, 1968; Calvert,
1969). As Calvert (1969) has pointed out, the problem concerns the post-
initiation stage of development of the flower buds since plants which fail to
produce a normal inflorescence always reveal a small immature truss which
invariably aborts.
Although there is much evidence on the importance of light conditions
and temperature in the control of the development of the tomato inflores-
cence (Lewis, 1953; Calvert, 1959, 1964; Kristoffersen, 1963; Lake, 1967;
Hurd and Cooper, 1967, 1970), there is little information available on the
critical levels of these environmental factors inducing abortion, and on the
stage of development during which the plant is most sensitive to adverse con-
16

ditions. It appears however, that the sensitivity is maximal during the late
stages of development of the inflorescence since abortion occurs when high
temperature and low light are applied at this time (Calvert, 1969).
Abortion is a c o m m o n and critical problem in winter-grown tomatoes in
Belgium and it is essential to try to avoid it. Because increasing light intensity
in glasshouses with artificial light would be t o o expensive, it is necessary to
find other means to prevent abortion. This is the objective of the present
research.
A better knowledge of the conditions giving rise to abortion is, however,
required and is a prerequisite for further studies. That is why in this first
paper, our aim is to gain accurate information on light conditions leading to
abortion. What is the effect of light intensity and photoperiod? At what stage
in the life of the plant do unfavourable light conditions mainly affect the
development of the truss? In order to avoid temperature effects, all the present
experiments were performed at the same constant temperature. The cultivar
'King Plus', c o m m o n l y grown in Belgium, was used in this study.

MATERIAL AND METHODS

G r o w t h c o n d i t i o n s . -- All the experiments were carried out in the growth

rooms of the p h y t o t r o n of the Botanical Department at Liege. Seeds o f the
cultivar 'King Plus' {Etabl. Pannevis, Duffel, Belgium) were germinated at
26°C in a peat c o m p o s t (TKS1 from Floratorf, Oldenburg, Germany). After
2 weeks, plants were pricked o u t in 7 cm pots filled with TKS: peat com-
post (Floratorf) and at an appropriate time into 14 cm pots. During growth,
the day and night temperature was maintained at a constant 20 ° C. We chose
this high temperature as it is known that a higher temperature is less favourable
to inflorescence development. Light was provided exclusively b y white flu-
orescent tubes (ACEC LF-40W/2 4 3 0 0 ° K or P h y t o r C.R.H.Lg.). The light
intensities used were low, which was also in order to obtain high percentages
of abortion.
There were 12 plants per treatment and at least one replication. In most
treatments, the data were recorded in respect of the first and second in-
florescences.

R e c o r d i n g m e t h o d s . - - Flower-initiation, development of the inflorescence and

plant growth were considered. For practical reasons the observations had to
be stopped after flower formation, so that no data on the fruits are available.
These will be collected in the near future.
For flower-initiation, the data presented refer to (a) the n u m b e r of days
from sowing to the macroscopic appearance of the truss, recorded when the
sepals of the first flower could be visualized individually; (b) the number of
leaves formed before inflorescence initiation.
For the development of the inflorescence, the data refer to (a) the n u m b e r
of days from the appearance of the truss to the first anthesis; (b) the per-
 17

centage of plants flowering at the considered truss using the anthesis of at

least 1 flower as the criterion; the complete failure of the truss to develop
any flower bud to the stage o f anthesis indicates the abortion of the inflores-
cence.
Data regarding plant growth refer to (a) the stem length measured from
the cotyledonary node to the shoot tip; (b) the rate of leaf production
determined b y recording periodically all the leaves having a blade more than
1 cm long.

RESULTS

Effects o f photoperiod and light intensity on flower-initiation, development

o f the inflorescence and plant growth. -- After pricking-out, plants were
grown in 4 light regimes. There were 2 photoperiods -- 8 and 16 h -- and 2
light intensities -- 9,000 ergs cm -2 s'' and 18,000 ergs cm -2 s" at the top of
the plants.
In these different conditions, the flowering-responses for the first 2 in-
florescences are shown in Table 1. The appearance of the inflorescences is
earlier and occurs after the formation of fewer leaves, in days with higher
light flux integrals. However, in days with similar light flux integrals, flow-
ering initiation is hastened in short days.

TABLE 1

E f f e c t o f p h o t o p e r i o d a n d light i n t e n s i t y o n f l o w e r - i n i t i a t i o n , i n f l o r e s c e n c e d e v e l o p m e n t
and g r o w t h . LI : L o w light i n t e n s i t y ; HI : High light i n t e n s i t y . Means a n d p e r c e n t a g e s w i t h
95% c o n f i d e n c e limits. F o r p e r c e n t a g e s , c o n f i d e n c e limits in square b r a c k e t s are read
directly f r o m P e a r s o n and H a r t l e y (1969, see Table 41 ).

Light regime

8h-LI 8h-HI 16h-LI 16h-HI

N u m b e r o f days
f r o m sowing t o 1st truss 78.1 +- 5.69 48.5 +- 1.00 52.1 ± 1.37 40.6 +-0.94
m a c r o s c o p i c ap- 2nd truss > 92 61.1 +- 1.58 66.5 +- 2.18 51.3 +- 1.09
pearance of the
N u m b e r o f leaves
formed before 1st truss 12.2+_0.65 9.2_+0.27 9.8+-0.25 9.3+-0.34
initiation o f t h e 2nd truss > 17 13.2 +-0.48 14.1 -+0.30 12.7 +-0.57

P e r c e n t o f flow- 1st truss 0 [0--14] 62.5 [ 4 1 - - 8 1 ] 0 [0--14] 100 [ 8 6 - - 1 0 0 ]

ering plants at
the 2nd truss 0 [0--14] 100 [ 8 6 - - 1 0 0 ] 18.2 [ 6 - - 3 9 ] 100 [ 8 6 - - 1 0 0 ]

N u m b e r o f days 1st truss -- 18.0 +- 1.51 -- 13.1 +-0.56

from macroscopic
a p p e a r a n c e to firSton d -- 15.8 +- 0.66 16.0 ± 3.90 13.0 +-0.48
truss
anthesis at t h e

S t e m length 51st day 56 ± 12.0 114 ± 12.4 156 ± 17.6 173 ± 7.8
( m m ) at t h e
 18

Regarding t h e d e v e l o p m e n t o f the i n f l o r e s c e n c e , it is clear t h a t it is mar-

k e d l y a f f e c t e d b y the daily light e n e r g y received. All plants f l o w e r e d at the
first 2 inflorescences in days with t h e higher light integral; all a b o r t e d in days
with the l o w e r light supply. O f the 2 t r e a t m e n t s with the same light integral,
i n f l o r e s c e n c e d e v e l o p m e n t is far b e t t e r in s h o r t days with high light i n t e n s i t y
t h a n in l o n g p h o t o p e r i o d s with l o w light intensity. L o o k i n g at t h e t i m e inter-
val f r o m t h e m a c r o s c o p i c a p p e a r a n c e o f t h e i n f l o r e s c e n c e t o t h e first anthesis,
it appears t h a t it is s h o r t e r w h e n the daily light s u p p l y is higher and, as s h o w n
at the s e c o n d i n f l o r e s c e n c e , it is identical in b o t h t r e a t m e n t s with t h e same
light flux integral.
S t e m g r o w t h is also a f f e c t e d b y t h e light t r e a t m e n t s . T h e higher the light
flux integrals, t h e taller t h e plants; w h e n t h e daily light s u p p l y is t h e same, t h e
stem g r o w t h is faster in long p h o t o p e r i o d s .
Figure 1 shows regression e q u a t i o n s f o r leaf n u m b e r o n days f r o m sowing

14,

/
12-

lO-

¢u
•
// ~ 9""
/o

¢.

e,

O" /~ 16h-LZ
• Bh -H T
2- O 8h-LI
/
o 2~ 3'1 3~ 4~ 4~ s'~ s'6

Days from sowing

Fig.1. E f f e c t o f l i g h t o n r a t e o f l e a f p r o d u c t i o n . H I : H i g h I n t e n s i t y - - LI : L o w I n t e n s i t y .
16 h -- high intensity : y = 0.34 x--4.42, r = 0.999;
16 h -- low intensity : y = 0.29 x --4.04, r = 0.998;
8 h -- high intensity : y = 0.31 x --4.91, r = 0.999;
8 h -- low intensity : y = 0.17 x--1.65, r = 0.999.
 19

and the respective regression lines. The rate at which new leaves are produced
is constant with time when the environmental factors remain constant; it in-
creases when the total a m o u n t of light energy supplied daily to the plants in-
creases and it is almost similar for plants grown in days with similar light in-
tegrals.

Effects o f transfers from favourable to adverse conditions on the develop-

m e n t o f the inflorescence.
Time o f effectiveness. -- Plants were grown from sowing in light conditions
allowing the development of the inflorescences in a high percentage of plants.
These so-called "favourable conditions" were daylength 12 h and light intensity
20,000 ergs cm -2 s-~ . In order to determine at what time during the life of
these plants insufficient light conditions are most effective in inducing the
abortion of the inflorescence, plants were transferred at different time inter-
vals, for 10 days, to adverse conditions: daylength 8 h; light intensity 12,000
ergs cm -2 s-~ . All the inflorescences from plants continuously kept in these
adverse conditions aborted.
The effect of these transfers on the development of the inflorescences is
shown in Figs 2 and 3. Giving the adverse light conditions before the 46th
day does n o t reduce the flowering-response at the first truss. From the 46th

70- Macroscopic appearance

~ of the truss

60-

= Control in favourabie conditions

~" 50-

°-
it-

l,- 40-

I:: 30"
k.
0

o
~" 20-

0. 1O-
N
Control in
adverse conditions

- ~e 3~ 4~ s~ ~6 7~

Transfer to adverse conditions

(days from sowing)

Fig.2. Effect of a transfer from favourable to adverse light c o n d i t i o n s , at different times

after sowing, on flowering of the first truss. The duration of the transfer was 10 days.
20

Macroscopic
appearance
~of the truss

90,
I!
Control in favourable conditions

80-

70-

~* 60-
¢
0
¢)

50-
o

40.
P
i.

0 30.

¢
~ 20.
0.

N
10-

C o n t r o l in adverse conditions

l_
o 2'6 3'~ ,~ s~ 6'6 7~
Transfer to adverse conditions (days from sowing)

Fig.3. E f f e c t o f a t r a n s f e r f r o m f a v o u r a b l e t o adverse l i g h t c o n d i t i o n s at d i f f e r e n t t i m e s
a f t e r sowing, o n f l o w e r i n g o f t h e s e c o n d truss. T h e d u r a t i o n o f t h e t r a n s f e r was 10 days.

day, i.e. the time of macroscopic appearance of the first truss, to the 56th day
adverse light conditions completely prevent inflorescence development.
Later, their effect is less and less marked with time (Fig.2).
For the second truss, similar results were obtained. However, the time of
greatest effectiveness is 10 days later than for the first truss; it corresponds
with the time of macroscopic appearance of the second truss (Fig.3).

D u r a t i o n o f t h e transfer. - - Plants grown until macroscopic appearance of

the first truss in favourable conditions were transferred for 2, 4, 6, 8 or 10
days to adverse conditions and then returned to favourable conditions. Re-
sults are summarized in Fig.4. They show that the n u m b e r of plants flow-
ering at the first truss decreases as the duration of the stay in adverse light
conditions increases. The effect of these adverse conditions is already detect-
able after 2 days and the abortion is almost complete after 10 days.
 21

80¸

60.

40-

e-
0

.? 20-

C \
D.
Control ir, adverse conditions ~ •
o

- g ~ ~ ; ~ ib

D u r a t i o n of thb s t a y in a d v e r s e
conditions (days)
Fig.4. Effect of the duration of the transfer from favourable to adverse light conditions
on flowering of the first truss. The transfer begins at the time of the macroscopic ap-
pearance of the truss.

Effect o f transfers from adverse to favourable conditions on the development

of the inflorescence.

Time o f effectiveness. -- Plants grown since sowing in adverse light condi-

tions were transferred at different time intervals to favourable conditions.
The duration o f the transfer was 15 days. As shown in Fig.5, favourable con-
ditions c a n n o t prevent inflorescence abortion when given before the 56th
or after the 71st day. When the plants were transferred at the time o f the
macroscopic appearance o f the inflorescence, i.e. at the 56th day, the favour-
able light conditions p r o m o t e flowering.
Later experiments have shown t hat t he prom ot i ve effect of the transfer
to favourable conditions was greater when it began a few days before the
macroscopic appearance o f the inflorescence. Similar results were obtained
at the second truss (Fig.6).

Duration o f the transfer. -- Plants grown in adverse light conditions were

transferred to favourable conditions just a few days (2--4) before t he macro-
scopic appearance of the first truss. The duration of t he transfer varied from
5 t o 20 days. The results in Fig.7 show t hat a stay of at least 15 days is re-
quired in o r d er to achieve the d e v e l o p m e n t o f the inflorescence in a n u m b e r
o f plants as high as in the controls c o n t i n u o u s l y kept in favourable light
22

Control in favourable conditions

70'

Macroscopic appearance
60" of the truss

.,i
50"

40'

0') 30"
¢:
i.
¢P
i=
0
q. 20-

e.

I:. 10-
Control in
~caodnV~irtSens
i

L_
o 4~ 5~ ~ ~6 i~1
Transfer to favourable conditions
(days from sowing)

Fig.5. Effect of a transfer from adverse to favourable light conditions, at different times
after sowing, on flowering of the first truss. The duration of the transfer was 15 days.

conditions; abortion is almost complete when the transfer is only for 10 days
or less.

DISCUSSION

The effect of environmental factors on the flowering of tomato have been

intensively studied. From the work already published, it appears that the ef-
fect of the daylength is small and hardly demonstrable because several other
factors influence the flowering-response of this species. For instance, it is
known that light intensity, temperature and nitrogen content in the soil (cf.
Wittwer and Aung, 1969) greatly affect flower-initiation. This complicated
situation can account for the different photoperiodic responses which were
successively attributed to tomato. From our experiments, using different
daylengths with a constant daily light energy supply, we can conclude, in
agreement with Wittwer (1963), Binchy and Morgan (1970), and Hurd
(1973), that tomato is a quantitative short-day plant: its flowering is earlier
and occurs after fewer leaves in short photoperiods than in long ones. In other
respects, our results clearly show that light intensity has a large effect on the
initiation of flowering in tomato, so that, irrespective of photoperiod, days
with higher light flux integrals are more suitable.
 23

90"
Control in favourable conditions

81:1,

70'
Macroscopic
appearance ~
of the truss
60

"0
e" 50-
0

o
J: 40-

C
':" 30-

2
~ 20-
C

~ 10-
Control in
~ caodnV~irt::ns
O-

L
o 4~ 56 7'1 86 101

Transfer to favourable conditions

(days from sowing)

Fig.6. Effect of a transfer from adverse to favourable light conditions, at different times
after sowing, on flowering of the second truss. The duration of the transfer was 15 days.

Inflorescence development in tomato is affected in the same way as the

initiation of flowering by light conditions. As in a number of other species
(cf. Nitsch, 1965) these two developmental processes thus have similar light
requirements. Inflorescence development is nevertheless more dependent on
short photoperiods and high light intensities than flower-initiation; in days
with similar light flux integrals anthesis occurs in almost all plants grown in
short days with a high intensity when abortion is almost complete in long
days with a low intensity.
Several studies demonstrated a flowering-inhibition by young leaves in
tomato (De Zeeuw, 1954; Leopold and Lam, 1960; Hussey, 1963} and it was
suggested that flower abortion is due to competition for available assimilates
between vegetative growth and inflorescence development in insufficient light
conditions (Hussey, 1963; Calvert, 1965, 1969; Cooper and Hurd, 1968;
Hand and Postlethwaite, 1971}.
Postulating a priority of vegetative over generative growth in the use of
24

Control in favourable
conditions f~)
80-

~ 60'

~ 40'

¢
L.
41
Q
~e. 20"

~ o.
1__

D u r a t i o n of t h e s t a y in f a v o u r a b l e
conditions (days)

Fig.7. Effect of the duration of the transfer from adverse to favourable light conditions
on flowering of the first truss. The transfer begins 2--4 days before the macroscopic ap-
pearance of the truss.

photosynthetic products under conditions of limited photosynthesis (Cooper,

1971) could explain why the effectiveness of a transfer from favourable to
adverse light conditions is greater than an inverse transfer (in our experi-
ments, complete abortion occurred after 10 days in insufficient light when
at least 15 days in favourable conditions were required in order to achieve
the development of the inflorescence). It might be, indeed, that during a
transfer to adverse conditions, the depletion in photosynthates first affects
the inflorescence; on the other hand, the increase in photosynthates after a
transfer to favourable conditions first favours vegetative growth (probably
the growth of y o u n g leaves). This hypothesis could also explain why a trans-
fer from adverse to favourable conditions is more effective in the prevention
of truss abortion when it begins some days before the macroscopic appearance
of the truss, these days being devoted to the p r o m o t i o n of vegetative growth.
Our experiments show that adequate light conditions are required particu-
larly at the time of, and just after, the macroscopic appearance of the truss,
i.e. during the late stage of the development of the inflorescence including
the lengthening of the peduncle and the growth of the floral organs. Calvert
(1969) came to the same conclusion. In his study, he showed that in contrast
with the first inflorescence, the second truss was not affected by adverse
light conditions. In our experiments, both inflorescences reacted similarly,
 25

suggesting that our observations are of general occurrence whichever the

truss studied. However, it must be pointed o u t that after a transfer to ad-
verse light conditions, flowering-inhibition at the second truss is of shorter
duration than at the first truss, while an inverse situation exists for flowering-
p r o m o t i o n after a transfer to favourable conditions. This p h e n o m e n o n could
be due to the fact that there are a greater n u m b e r of mature leaves supporting
the supply of photosynthates during the development of the second inflores-
cence than during the development of the first one. As a result, the effect of
adverse conditions would be more injurious to the first truss than to the
second one and favourable conditions would favour the second rather than
the first inflorescence. This fact, connected with the weak flowering-inhibi-
tion that Calvert obtained at the first truss could explain w h~ this author
did n o t observe any effect of adverse conditions on the development of the
second truss. So the discrepancy between the results o f the British worker
and our observations would be greater in semblance than in reality.
The fact that a transfer from adverse to favourable light conditions can
prevent abortion when given at a critical time during the life of the plant b u t
n o t later, suggests that there is a point of no return in the progress o f the
inflorescence to abortion. A similar conclusion can be drawn from experi-
ments using transfers from favourable to adverse light conditions: during the
normal development of the inflorescence to flower anthesis a point of no
return is likewise reached. A precise knowledge of the time after which abor-
tion is no longer possible could have value in practical applications.

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26

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