Growing Greenhouse Tomato and Sweet Pepper under Supplemental

Lighting: Optimal Photoperiod, Negative Effects of Long Photoperiod

and Their Causes
D.A. Demers A. Gosselin
Greenhouse and Processing Crops Centre de Recherche en Horticulture
Research Centre Université Laval
Agriculture and Agri-Food Canada Québec, QC, Canada, G1K 7P4
Harrow, On., Canada, N0R 1G0 Andre.Gosselin@fsaa.ulaval.ca

Keywords: Lycopersicon esculentum, Capsicum annuum, chlorosis, carbon metabolism,

light quality, pigment

Abstract
This paper reviews the available information on the photoperiod aspects of
the use of supplemental lighting for greenhouse tomato and sweet pepper
production. Optimal growth and yields of tomato and sweet pepper were obtained
under photoperiods of 14 and 20 hours, respectively. Longer photoperiods did not
further improve growth and yields and even decreased growth and yields in some
cases. Although long term use of continuous light is detrimental to tomato and
pepper plants, vegetative growth and fruit production of both species can be
improved by short term use (5 to 7 weeks) of continuous lighting. Compared to
shorter photoperiods, continuous light (24-h photoperiod) increased the leaf levels of
hexoses in tomato, of sucrose in pepper and of starch in both species. The
accumulation of starch and sugar in leaves under continuous light indicate a
limitation of tomato and pepper plants to export the photosynthate out of their
leaves. Such a limitation would explain the fact that extra light energy provided by
continuous lighting did not result into growth and yield gains. The increased leaf
hexose levels in tomato and increased leaf sucrose in pepper suggest that the limiting
steps of the export of photosynthate are respectively the synthesis of sucrose and the
loading of sucrose in the phloem. Under greenhouse conditions, continuous light
caused leaf chlorosis in tomato but not in sweet pepper. Development of leaf
chlorosis in tomato under continuous light was related to a decrease of the
chlorophyll concentration in the leaves. Compared to tomato, higher levels of
carotene and xanthophylls (photoprotective pigments) in pepper leaves probably
provided a better protection of the photosynthetic apparatus against excessive light,
thus preventing the destruction of chlorophylls and the development of leaf chlorosis
in pepper. The severity of leaf chlorosis varied with the type of lamps (high pressure
sodium, HPS versus metal halide, MH) used to provide the supplemental light,
indicating that the spectral composition of the light received by plants may also play
a role in the development of leaf chlorosis. Under continuous light, the response of
tomato and pepper plants to HPS lamps versus MH in the greenhouse differed from
the response in growth chambers. These differences between greenhouse and growth
chamber could be related to the light spectral quality (presence or absence of
natural light) and/or the daily variation in the climatic conditions (larger day/night
differential in greenhouse).
INTRODUCTION
The low natural light level during wintertime is the most important factor limiting
greenhouse vegetable productions in northern regions such as Canada. Many studies have
shown that using supplemental light during wintertime increased growth and yield of
greenhouse tomato (Lycopersicon esculentum Mill.) (Dorais et al., 1988; Vézina et al.,
1991), cucumber (Cucumis sativus L.) (Blain et al., 1987; Turcotte and Gosselin, 1988),
sweet pepper (Capsicum annuum L.) (Demers et al., 1991) and lettuce (Lactuca sativa L.)

Proc. 4th IS on Artif. Light.

Ed. M. Dorias 83
Acta Hort. 580, ISHS 2002
(Gaudreau et al., 1994). Most studies investigated the effects of supplemental light levels
on plants while only a few studied the photoperiod effects. It has been known for a long
time that exposing tomato plants to continuous light (24-h photoperiod) causes the
development of leaf chlorosis and growth reductions (Guthrie, 1929; Kristoffersen, 1963).
However, the cause of these problems remains unclear. A short review on the factors
possibly involved in the development of the negative effects of long photoperiods on
tomato and sweet pepper plants was published by Demers et al. (1999). This paper
presents a discussion on the photoperiod aspects (optimal photoperiods, negative effects
of longer photoperiods and their causes) of supplemental lighting for greenhouse tomato
and sweet pepper production.

OPTIMAL PHOTOPERIODS
For tomato, best growth and yield were obtained under a photoperiod of 14 hours
(Vézina et al., 1991; Demers et al., 1998b). Photoperiods longer than 14 h did not further
increase yield. Photoperiods of 20 and 24 h can even decrease yield and caused leaf
chlorosis (after 6 to 8 weeks) (Vézina et al., 1991; Demers et al., 1998b). Although long
term use of a 17-h photoperiod does not increase growth and yield compared to 14 h, it
might be interesting to extend the photoperiod to 17 h in order to increase total light
provided to plants especially during the months with the lowest natural light levels
(December-January). However, if a 17-h photoperiod is used, it is important that the dark
period be uninterrupted, since splitting the dark period of 7 h in two short nights of 3.5 h
(separated by a light period of 4 h) caused leaf chlorosis and decreased growth and yield
(Vézina et al., 1991).
For sweet pepper, a 20 h-photoperiod was optimal for plant growth and
productivity (Demers et al., 1998a). Yield under continuous light (24-h photoperiod) was
equivalent to yield under photoperiods of 15 or 16 h (Costes et al., 1970; Demers et al.,
1998a). Extension of the photoperiod from 15 or 16 h to 24 h decreased the average size
of pepper fruits (Costes et al., 1970; Demers et al., 1998a). Continuous light caused some
leaf deformities (wrinkles) but no chlorosis in sweet pepper grown in greenhouses.
Although long term use of continuous light is detrimental to tomato and pepper
plants, tomato and sweet pepper plants can take advantage of the extra light energy
provided by continuous lighting for a short period of time. Early vegetative growth and
fruit production of tomato and pepper plants were generally improved under continuous
light compared the 14-h photoperiod (Demers et al., 1998a, 1998b). However, after that
initial period, plants under continuous light grew more slowly than plants exposed to 14-h
photoperiod; so that tomato and pepper plant growth and yield under 14-h photoperiod
were then equal to or higher than under continuous light at the end of the experiment.
Costes et al. (1970) also observed that continuous light improved the early performance
(hastening of flowering and fruit set, increased early yield) of sweet pepper plants
compared to a 15-h photoperiod. Therefore, it might be possible to use continuous light
for a short period of time (5 to 7 weeks) to improve growth of tomato and sweet pepper,
especially during the months with the lowest natural light levels (December and January).
However, such a practice should be investigated in order to determine if short term use of
continuous light might have residual negative effects on tomato and sweet pepper plants.
NEGATIVE EFFECTS OF LONG PHOTOPERIODS AND THE FACTORS
INVOLVED IN THEIR DEVELOPMENT
Tomato and sweet pepper plants do not take advantage (no increase in yield) when
grown under photoperiods longer than 14 h (tomato) or 20 h (pepper). Tomato plants, but
not sweet pepper, develop leaf chlorosis under continuous light. In the next sections, we
will examine the role of the carbon metabolism, pigments, light spectral quality and
day/night temperature differential in the development of these negative effects of long
photoperiods.

84
Carbon Metabolism
High starch and soluble sugar accumulations were observed in leaves of tomato
plants grown under long photoperiods, and it was suggested that these accumulations
could be related to the development of the leaf chlorosis (Bradley et al., 1985; Logendra
et al., 1990; Dorais, 1992). Studies on other species support the hypothesis of a
relationship between leaf chlorosis development and starch and sugar accumulations. For
example, continuous light caused increased leaf starch and hexose accumulations and leaf
chlorosis of eggplants (Solanum melongena L.) (Murage et al., 1996). However,
eggplants growing under continuous light but in a CO2-free atmosphere for 12 h per day
accumulated less starch and hexoses, and did not develop leaf chlorosis.
Exposure of tomato and sweet pepper plants to continuous light resulted in
increased foliar contents in starch in tomato and sweet pepper, in hexoses (glucose and
fructose) in tomato and sucrose in sweet pepper (Dorais et al., 1996; Demers et al., 1998a,
1998b). However, the reduction of the number of fruits on the plants did not modify the
pattern of accumulation of starch and sugars in leaves of tomato and sweet pepper plants
exposed to photoperiods of 14 and 24 h (Demers et al., 1998a, 1998b). Moreover, the
reduction of the number of fruits on the plants did not influence the severity nor the date
of appearance of the foliar chlorosis in tomato plants grown under continuous light. This
indicates that accumulations of starch and soluble sugars are not caused by a limiting sink
capacity. If there is a relationship between the excessive starch and soluble sugar
accumulations and the development of the negative effects (leaf chlorosis, decreased
growth and productivity) of the long photoperiods on tomato and sweet pepper, it is most
likely a limitation of the carbon metabolism at the leaf level which is responsible for these
accumulations.
In tomato, the use of continuous light caused, in addition to the foliar chlorosis
and increased foliar contents in starch and hexoses, a reduction of the photosynthesis rate
and of the activity of the sucrose phosphate synthase (SPS) enzyme (Demers, 1998).
These reductions in photosynthesis and of SPS activity occurred between 6th and 8th week
under continuous light, i.e. about at the same time as the foliar chlorosis appeared, while
starch and hexoses contents in leaves increased during the first 4 weeks of the experiment.
Since the reduction of the SPS activity occurred after the increase in starch and hexoses, it
is thus impossible that the reduction of the SPS activity is responsible for these
accumulations. However, it is possible that the SPS activity in vivo is limiting, which
would explain the hexose increase. This suggests the limiting step of the export of
photosynthates is the synthesis of sucrose in tomato and would explain the absence of
growth and the productivity increase under continuous light. Furthermore, the increased
hexose levels in the cytoplasm, by a feedback effect, would limit the export of the triose-
phosphate (photosynthesis products) out of the chloroplast, which would then be
redirected towards starch synthesis, thus explaining the increased starch contents.
Moreover, the increased accumulation of starch would generate, by a feedback effect, an
overload of the Calvin cycle, which would gradually cause the observed decrease of the
CO2 fixation rate. Are the starch accumulations responsible for the leaf chlorosis in
tomato? It is possible that the overload imposed on the Calvin cycle (decreased
photosynthesis) could limit the use of the reducing potential (ATP, NADPH) produced by
the luminous phase of photosynthesis, thus causing an overload on the electron transport
chain and the photo-oxidation of the chlorophylls (decrease in the leaf chlorophyll
contents), and thus explaining the observed leaf foliar chlorosis. Transgenic tomato plants
(in which a gene coding for the SPS enzyme was incorporated and overexpress this
enzyme) could be used in future studies to test if accumulations of starch in leaves are
responsible for the development of chlorosis observed in tomato plants exposed to
continuous light. Transgenic tomato plants (overexpressing SPS) have higher
photosynthesis rates and accumulate less starch and more sucrose than non-transformed
plants, especially under conditions of saturating light and CO2 (Galtier et al., 1993, 1995;
Micallef et al., 1995). One can put forth the assumption that, under continuous light, leaf
starch contents would be lower in transgenic plants than in normal plants. If this is the

85
case, the reduction of the leaf starch content in transgenic plants should thus prevent the
development of the leaf chlorosis, or at least decrease its severity.
In sweet pepper, the use of continuous light caused an increase in the leaf starch
and sucrose contents, but did not affect leaf hexose contents, photosynthesis rates and
SPS activity (Demers, 1998). The increased foliar contents in sucrose indicate that SPS
activity in sweet pepper is not limiting as in tomato. Increased accumulation of starch in
sweet pepper plants exposed to continuous light would be explained by the fact that
continuous light results in a longer period of time over which starch synthesis occur, but
without overloading the starch synthesis pathway. Thus, starch accumulation in sweet
pepper under continuous light would not be important enough to cause a reduction in CO2
fixation (no overload of the Calvin cycle). Increased leaf contents in sucrose suggest that
sucrose export would be possibly limiting. In sweet pepper plants, the export rate of
carbon (as sucrose) out of the leaf is constant, and the export rate would be limited at the
level of the loading of sucrose in the phloem (Grange, 1985, 1987). This would explain
why the growth and the productivity of the sweet pepper plants do not increase under
continuous light.

Pigments
In growth chambers, continuous light caused leaf chlorosis, decreased
photosynthesis rates, and reductions in leaf contents in pigments (chlorophyll a and b,
carotene, xanthophylls) in both tomato and sweet pepper plants (Demers, 1998). Leaf
chlorosis, decreased photosynthesis rates and loss of pigments were more important and
occurred earlier in tomato plants than in sweet pepper. Compared to sweet pepper plants,
EPS ratio (epoxidation state of the pigments of the xanthophyll cycle) was lower in
tomato, indicating a greater need for energy dissipation and a more important state of
stress (caused by excessive light). Pigments such as carotene and xanthophylls
(violaxanthin, antheraxanthin, zeaxanthin) play a significant role in the protection of the
photosynthetic apparatus against damage that could be caused by an excess of light.
Carotene and xanthophyll levels were higher in sweet pepper plants than in tomato. Thus,
sweet pepper has a better protection against the degradation of chlorophylls, which would
explain why leaf chlorosis appeared later and were less severe in sweet pepper.

Light Spectral Quality

In tomato plants grown under continuous light in greenhouses, leaf chlorosis were
slightly more severe and photosynthesis rate decrease more important when the artificial
light was provided by metal halide (MH) lamps compared to high pressure sodium (HPS)
lamps (Demers, 1998). Differences in the severity of the foliar chlorosis were observed
depending on the type of lamps used to grow eggplants under continuous light (Murage et
al., 1997). This indicates that the spectral quality of the light provided to plants (by
different lamps) could also play a role in the development of the negative effects of long
photoperiods. Compared to MH lamps, HPS lamps give less light energy between 400
and 500 nm (blue light), and more light energy between 600 and 700 nm (light red),
which could be related to the difference in the response of tomato plants to the two types
of lamps.
In tomato grown in growth chambers, leaf chlorosis, decreased photosynthesis
rates, and reductions in leaf contents in pigments caused by continuous light were more
important when light was provided by HPS lamps compared to MH lamps (Demers,
1998). This differs from results in greenhouses where leaf chlorosis and decreased
photosynthesis rates were more important under MH lamps than HPS. Moreover, leaf
chlorosis were observed on sweet pepper plants grown in growth chambers, but not on
those grown in greenhouses. The presence of natural light in the greenhouse, but not in
growth chambers, may explain the different results between the two experiments. In order
to determine why the response of tomato plants to lamps HPS and MH is modified by the
presence or the absence of natural light, new studies on the influence of the spectral
quality of the light (various sources of light, with or without natural light) provided to the

86
plants should be undertaken.
Day/Night Temperature Differential
Another factor that may explain the different results between the greenhouse and
growth chamber experiments is the greatest variation of the environmental conditions in
greenhouse (influenced by the external conditions) compared with the conditions in
growth chambers (humidity and light constant, small temperature variation (< 3oC). Leaf
chlorosis were also observed on sweet pepper plants grown in growth chambers under a
continuous lighting and a constant temperature (Nilwik, 1981). Kristoffersen (1963) and
Hillman (1956) observed that, under continuous lighting, leaf chlorosis developed on
tomato plants subjected to a constant temperature regime, but not on plants subjected to
variable temperature regime (day/night differential of 8-10oC). This could explain why
chlorosis were observed on sweet pepper plants grown under continuous lighting in
growth chambers but not on those grown in greenhouses. This could also explain why
chlorosis in tomato developed faster (2 to 4 weeks) and became much more severe (leaves
almost completely yellow or white) in growth chambers than in greenhouses (6-7 weeks
for chlorosis appearance). New research should be undertaken in order to look further into
the impact of temperature regimes (day/night differential) on photosynthesis, carbon
metabolism and leaf pigment contents of sweet pepper and tomato plants grown under
continuous light.

Literature Cited
Blain, J., Gosselin, A. and Trudel, M.J. 1987. Influence of HPS supplementary lighting on
growth and yield of greenhouse cucumbers. HortScience 22:36-38.
Bradley, F.M. and Janes, H.W. 1985. Carbon partitioning in tomato leaves exposed to
continuous light. Acta Hort. 174:293-302.
Costes, C. and Milhet, Y. 1970. Étude photophysiologique du poivron. Effets de la
lumière artificielle et du gaz carbonique sur la croissance et sur la production de fruits
sous serre. Lab Photophys, Dep Phys Veg INRA, CNRA, Route de St-Cyr 78,
Versailles, France. 45 p.
Demers, D.A. 1998. Physiologie, photosynthèse et métabolisme carboné de plants de
tomate (Lycopersicon esculentum Mill.) et de poivron (Capsicum annuum L.) cultivés
sous de longues photopériodes. Thèse de Doctorat, Faculté des Études Supérieures,
Université Laval, Ste-Foy, Québec, Canada.
Demers, D.A., Charbonneau, J. and Gosselin, A. 1991. Effets de l’éclairage d’appoint sur
la croissance et la productivité du poivron cultivé en serre. Can. J. Plant Sci. 71:587-
594.
Demers, D.A., Wien, H.C. and Gosselin, A. 1998a. Effects of supplemental light duration
on greenhouse sweet pepper plants and fruit yields. J. Amer. Soc. Hort. Sci. 123:202-
207.
Demers, D.A., Dorais, M., Wien, H.C. and Gosselin, A. 1998b. Effects of supplemental
light duration on greenhouse tomato (Lycopersicon esculentum Mill.) plants and fruit
yields. Scientia Hort. 74:295-306.
Demers, D.A. and Gosselin, A. 1999. Supplemental lighting of greenhouse vegetables:
Limitations and problems related to long photoperiods. Acta Hort. 481:469-473.
Dorais, M. 1992. Aspect culturaux et physiologiques de la tomate et du poivron de serre
soumis à un éclairage d’appoint. Thèse de Doctorat, Faculté des Études Supérieures,
Université Laval, Ste-Foy, Québec, Canada.
Dorais, M., Gosselin, A. and Trudel, M.J. 1988. Growing systems for greenhouse tomato
production under high photosynthetic photon flux density supplemental lighting. Acta
Hort. 230:405-412.
Dorais, M., Yelle, S. and Gosselin, A. 1996. Influence of extended photoperiod on
photosynthate partitioning and export in tomato and pepper plants. New Zealand J.
Crop Hort. Sci. 24:29-37.
Galtier, N., Foyer, C.H., Huber, J., Voelker, T.A. and Huber, SC. 1993. Effects of

87
 elevated sucrose-phosphate synthase activity on photosynthesis and assimilate
partitioning and growth in tomato (Lycopersicon esculentum var UC82B). Plant
Physiol. 101:535-543.
Galtier, N., Foyer, C.H., Murchie, E., Alred, R., Quick, P., Voelker, T.A., Thépenier, C.,
Lascève, G. and Betsche, T. 1995. Effects of light and atmospheric carbon dioxide
enrichment on photosynthesis and carbon partitioning in the leaves of tomato
(Lycopersicon esculentum L.) plants over-expressing sucrose phosphate synthase. J.
Exp. Bot. 46:1335-1344.
Gaudreau, L., Charbonneau, J., Vezina, L.P. and Gosselin, A. 1994. Photoperiod and PPF
influence growth and quality of greenhouse-grown lettuce. HortScience.
29:1285-1289.
Grange, R.I. 1985. Carbon partitioning and export in mature leaves of pepper (Capsicum
annuum). J. Exp. Bot. 36:734-744.
Grange, R.I., 1987. Carbon partitioning in mature leaves of pepper: Effects of transfer to
high or low irradiance. J. Exp. Bot. 38:77-83.
Guthrie, J.D. 1929. Effects of environemental conditions on chloroplast pigments.
Contrib. Boyce Thompson Inst. 2:220-251.
Hillman, W.S. 1956. Injury of tomato plants by continuous light and unfavorable
photoperiodic cycles. Am. J. Bot. 43:89-96.
Kristoffersen, T. 1963. Interactions of photoperiod and temperature on growth and
development of young tomato plants. Physiol. Plant. Supplementum. 98 pp.
Logendra, S., Putman, J.D. and Janes, H.W. 1990. The influence of light period on carbon
partitioning, translocation and growth in tomato. Scientia Hort. 42:75-83.
Micallef, B.J., Haskins, K.A., Vanderveer, P.J., Roh, K.S., Shewmaker, C.K. and
Sharkey, T.D. 1995. Altered photosynthesis, flowering, and fruiting in transgenic
tomato plants that have increased capacity for sucrose synthesis. Planta 196:327-334.
Murage N.M., Watashiro N. and Masuda M. 1996. Leaf chlorosis and carbon metabolism
of eggplant in response to continuous light and carbon dioxide. Scientia Hort. 67:27-
37.
Murage, E.N., Sato, Y. and Masuda, M. 1997. Influence of light quality, PPFD and
temperature on leaf chlorosis of eggplants grown under continuous illumination.
Scientia Hort. 68:73-82.
Nilwik, H.J.M. 1981. Growth analysis of sweet pepper (Capsicum annuum L.) 1.
Influence of irradiance and temperature under glasshouse conditions in winter. Ann.
Bot. 48:129-136
Turcotte, G. and Gosselin, A. 1988. Influence du mode de distribution d'un éclairage
d'appoint et de la photopériode sur la croissance et les rendements du concombre de
serre. Can. J. Plant Sci. 68:535-542.
Vézina, F., Trudel, M.J. and Gosselin, A. 1991. Influence du mode d’utilisation de
l’éclairage d’appoint sur la productivité et la physiologie de la tomate de serre. Can. J.
Plant Sci. 71:923-932.

88