Amer J of Potato Res (2005) 82:353-367
Light E f f e c t s on t h e G r o w t h a n d M o r p h o g e n e s i s o f P o t a t o
(Solanum tuberosum) In Vitro: A R e v i e w
J a n e t E. A. S e a b r o o k
Potato Research Centre, Agriculture and Agri-FoodCanada, P.O. Box 20280, Fredericton, New Brunswick, E3B 4Z7 CANADA
Tel: (506) 452-4831;Email: seabrookj@em.agr.ca
ABSTRACT
Growth, m o r p h o g e n e s i s , a n d t u b e r i z a t i o n o f p o t a t o
t i s s u e s in vitro are affected by light. M e a s u r e m e n t s o f
the v a r i o u s a s p e c t s o f light t h a t c o n t r o l d e v e l o p m e n t
a n d growth o f p o t a t o are o u t l i n e d . Physical p a r a m e t e r s
like light sources, delivery o f t h e light source, a n d t h e
d e g r a d a t i o n of c u l t u r e m e d i a by light are discussed.
I r r a d i a n c e , p h o t o a u t o t r o p h i c growth in vitro, s p e c t r a l
wavelength, a n d p h o t o p e r i o d modify t h e r e s p o n s e s o f
p o t a t o t i s s u e s i n culture. A c c l i m a t i z a t i o n o f t i s s u e cul-
t u r e p l a n t l e t s , v e g e t a t i v e growth, a n d t h e p r o d u c t i o n ,
quality, a n d d o r n m a c y o f m i c r o t u b e r s a r e modified b y
light. New light s o u r c e s such as l i g h t - e m i t t i n g diode
( L E D ) l a m p s a r e b e c o m i n g a v a i l a b l e for in vitro
r e s e a r c h a n d for m i c r o p r o p a g a t i o n o f p o t a t o . P u l s e d o r
c h o p p e r light has t h e p o t e n t i a l t o save e n e r g y costs.
Light effects o n p o t a t o p r o t o p l a s t s , a n t h e r c u l t u r e , v i r u s
e r a d i c a t i o n , a n d in vitro c o n s e r v a t i o n are discussed.
P o t e n t i a l n e w r e s e a r c h a r e a s are the effect of t h e spec-
t r a l q u a l i t y of light o n r e g e n e r a t i o n o f s h o o t s a n d
somatic e m b r y o s in vitro, end-of-day red a n d far-red
light t r e a t m e n t s , axillary s h o o t f o r m a t i o n i n c u l t u r e d
p l a n t l e t s , a n d t h e u s e o f LEDs. T h e i n f l u e n c e o f
m o n o c h r o m a t i c s p e c t r a l f i l t e r s o n growth a n d develop-
m e n t o f p o t a t o e s i n t i s s u e c u l t u r e could p o t e n t i a l l y lead
t o i m p r o v e m e n t s i n p r o d u c t i v i t y . The r e l a t i o n s h i p
b e t w e e n daily q u a n t u m light i n t e g r a l a n d p h o t o p e r i o d
a n d t h e i r effects o n growth a n d m o r p h o g e n e s i s o f the
p o t a t o will provide some useful a r e a s o f research.
Accepted for publication 11 January 2005.
ADDITIONAL KEY WORDS: irradiance, filters, morphogenesis, pho-
toautotrophic, photoperiod, spectral quality,tuber
353
RESUMEN
Los p r o c e s o s de c r e c i m i e n t o , morfog~nesis y t u b e r -
izaci6n de los t e j i d o s de p a p a in vitro s o n afectados p o r
la luz. Mediciones de los varios a s p e c t o s de la luz que
c o n t r o l a n el c r e c i m i e n t o y d e s a r r o l l o de la p a p a s o n
delineadas. Parfimetros f[sicos t a l e s como la f u e n t e de
luz, difusi6n y su efecto en la d e g r a d a c i 6 n del medio de
cultivo s o n discutidos. La i r r a d i a c i 6 n , el c r e c i m i e n t o
f o t o - a u t o t r 6 f i c o in vitro, la l o n g i t u d de o n d a e s p e c t r a l y
el foto p e r i o d o modifican las r e s p u e s t a s de los t e j i d o s de
la p a p a e n cultivo. La a c l i m a t a c i 6 n de las plAntulas de
cultivo de tejidos, c r e c i m i e n t o v e g e t a t i v o y p r o d u c c i 6 n ,
calidad y d o r m a n c i a de los m i c r o t u b ~ r c u l o s s o n modifi-
cadas p o r la luz. Nuevas f u e n t e s de luz como a q u e l l a
e m i t i d a por las 1Amparas de diodo (LED) estAn s i e n d o
d i s p o n i b l e s p a r a la i n v e s t i g a c i 6 n in vitro y la micro-
p r o p a g a c i 6 n de la papa. La luz i n t e r m i t e n t e t i e n e el
p o t e n c i a l de a h o r r a r los costos de energ/a. Los efectos
de la luz sobre los p r o t o p l a s t o s de papa, cultivo de
a n t e r a s , e r r a d i c a c i 6 n de v i r u s y c o n s e r v a c i 6 n in vitro
s o n discutidos. Nuevas ~reas p o t e n c i a l e s de investi-
gaci6n son, el efecto de la calidad e s p e c t r a l de la luz
sobre la r e g e n e r a c i 6 n de los b r o t e s y los e m b r i o n e s
somfiticos in vitro, t r a t a m i e n t o s con luz r o j a del final del
dia y del lado l e j a n o del e s p e c t r o i n f r a r r o j o , f o r m a c i 6 n
a x i l a r de b r o t e s e n plAntulas c u l t i v a d a s in vitro y el uso
de LEDs. La i n f l u e n c i a de los f i l t r o s e s p e c t r a l e s
monocromfiticos s o b r e el c r e c i m i e n t o y d e s a r r o l l o de
plfintulas de p a p a e n cultivo de t e j i d o s p o d r i a p o t e n c i a l -
ABBREVIATIONS:CO2, carbon dioxide; B, blue; DOE diffusive optical
fibres; DLI,dailylight integral;DQLI,daily quantum light integral;FR, far
red; LAA,indole acetic acid; LEDs, light-emittingdiodes; PAR,photosyn-
thetically active radiation; PPF, photosynthetic photon flux; PPFD, pho-
tosynthetic photon flux density; R, red; UV, ultraviolet
354 AMERICAN JOURNAL O F POTATO RESEARCH
m e n t e conducir a u n m e j o r a m i e n t o e n la productividad.
La r e l a c i 6 n e n t r e la c a n t i d a d diaria de luz i n t e g r a l y f o t o
p e r i o d o y s u s e f e c t o s sobre el c r e c i m i e n t o y m o r f o g ~ n e -
sis de la p a p a proporcionarKu a l g u n a s ~weas d t i l e s de
investigaci6n.
INTRODUCTION
Potato ( S o l a n u m tuberosum L.) is a crop that grows best
in cool temperate climates with full sunlight, moderate day-
time temperatures, and cool nights. Short days generally
induce tubers in potatoes, although many modern cnltivars
can initiate tuberization in the long days of north temperate
regions (Hawkes 1992; Tam et al. 1992). Greenhouse-grown
potatoes require artificial light to extend the day during the
fall, winter, and spring seasons. In northern climates green-
house-grown plants may need supplemental light throughout
the day to produce a crop of minitubers.
The common perception that i n vitro cultures are less
sensitive to light than plants i n vivo is not the case (Aksenova
et al. 1989, 1994; Seabrook 1987; Seabrook et al. 1993). Pho-
toperiod, irradiance, and light spectral quality can be used to
control the growth and morphogenesis of potato tissues and
organs in vitro, thereby in some instances, avoiding the use of
growth regulators, which could possibly cause off-types
(Aksenova et al. 1989, 1994; Seabrook 1987; Seabrook et al.
1993; Seabrook and Douglass 1998; Wilson et al. 1993). Potato
plantlets i n vitro can produce microtubers in response to
inductive short photoperiods (Garner and Blake 1989; Pelacho
and Mingo-Castel 1991; Seabrook et al. 1993). Irradiance levels
influence the etiolation of potato plantlets i n vitro (Fujiwara
and Kozai 1995) and light quality can alter the morphology of
plantlets (Seabrook 1987; Seabrook and Douglass 1998; Wilson
et al. 1993).
Potato tissues in vitro are generally not autonomous for
photosynthesis and frequently depend on a source of organic
carbon such as sucrose (George 1986). Experimental regimes
for autotrophic growth of potato i n vitro using high levels of
irradiance and supplemental CO2 have been reported (Kozai et
al. 1988; Pruski et al. 2002). However, facilities using high irra-
diance levels are costly to build and require considerable
inputs of electrical energy to operate at optimal temperatures
(18 to 20 C) for potato tissue cultures.
Light is the electromagnetic radiation that causes photo-
chemical reactions in plants (Fujiwara and Kozai 1995; Tantau
Vol. 82
1997), and only a portion of this light is visible to the human
eye. Light requirements for photomorphogenesis are in the
near-nltra violet (300-380 nm), blue (430-490 nm), red (640-700
urn), and far-red (700-760 nm) regions of the light spectrum,
and for photosynthesis between 400 and 700 nm (Hart 1988).
Many fluorescent lamps used to illuminate plant tissue cul-
tures emit light deficient in the far-red wavelengths, thus care
must be taken to provide the spectral balance suitable for
growth of cultures (Seabrook 1987; v~rflson et al. 1993). Mor-
phogenesis of potato tissue cultures can be manipulated by
light regimes, and to improve efficiency of propagation (Kozai
et al. 1988; Niu et al. 1997; Prnski et al. 2002; Seabrook 1987;
Seabrook and Douglass 1998; Seabrook et al. 1993; Wilson et
al. 1993). The terms "daylength" or "photoperiod" are often
used to describe the alternating light and dark cycles within a
24-h period, but it is the length of the dark period that deter-
mines the photoperiodic response (Vince-Prue 1994).
This review will outline the importance of precisely defin-
ing the light regimes for potato tissue cultures used in research
and commercial production, outline alternative methods of
controlling growth and morphogenesis of potato tissues i n
vitro, and offer insights into possible productive areas of
future investigation.
Light Measurements
Irradiance, irradiant energy flux density (Wm-2), or inci-
dent light flux density (pmol'm-2s-1) (Fujiwara and Kozai
1995) are commonly called light intensity. Irradiance is mea-
sured in different ways: photometrically, radiometrically, and
in quantum terms, each of these components relates to a dif-
ferent property of radiant energy (Hart 1988). Light meters
usually have three different sensors that perceive these differ-
ent properties of radiant energy. Photometric measurements
(lumens = lux or foot candles), which measure brightness and
not energy or photons, are used by lighting engineers and in
the light meters in cameras (Hart 1988).
Photosynthetically active radiation or solar radiation on a
plane surface is the action spectrum used by plants for photo-
synthesis (ca. 350 to 730 um) (McCree 1972). Photosynthetic
research uses photosynthetic photon flux densities (PPFD) or
photosynthetic photon flux (PPF) to measure solar radiation
in watts m -2 (Sager and McFarlane 1997). Current horticultural
usage measures PPFD by using the pyranometer probe of a
light meter. Radiometric or pyranometric measurements of
light detect all forms of radiant energy including infra-red radi-
2005 SEABROOK: LIGHT AND POTATO I N VITRO
ation and are not suitable for morphogenetic research on
plants where comparisons of the energy associated with dif-
ferent wavelengths of light are important (Hart 1988). The
daily quantum light integral (DLQI) is the PPF incoming per
unit ground area, expressed as the sum of the total light for a
day (mol m -z"d 1 = MJm-2d-1) and has been useful in horticul-
tural research and pot plant production (White and Warrington
1984). Young plants appear to be particularly sensitive to DLQI
(White and Warrington 1984). The greenhouse industry has
been able to avoid the use of growth regulators (Carvalho and
HeuveIink 2001) by focussing on DLQI.
Research on plant morphogenesis requires the use of
quantum sensors that measure light in ~mol" m -2 s -1. Whimijan
and Heins (1983) provided empirical conversions for photo-
metric, radiometric, and quantum units as a guide for various
commonly used light sources in the scientific literature. Light-
unit conversions require information on lamp type (e.g., cool-
white, warm white, Gro-LtLx, etc.) used and the wavelength
interval emitted from the light source (Both 1994; Thimijan
and Heins 1983). Consequently, conversion from one unit of
light measurement to another is fraught with difficulty and can
only be estimated if the lamp type, age, and output are care-
fully defined. As this information is not available in the refer-
ences cited herein, no attempt has been made to convert units
for this review.
Spectroradiometers measure radiation at specific wave-
lengths of light and in plant science are usually calibrated to
measure at 10-nm intervals providing a graphical presentation
with quantity on the vertical axis and nanometers on the hori-
zontal axis.
Light Sources
Controlled environment facilities for plant tissue culture
use fluorescent lamps occasionally supplemented with incan-
descent lamps to provide additional red (R) portions of the
spectrum. The growth and morphogenesis of potato plantlets
in vitro can be radically altered by using different types of
lamps with varying spectral qualities (Charles et al. 1992;
Sarkar et al. 1996; Seabrook 1987; Wilson et al. 1993). Light
spectral distribution of fluorescent lamps can vary consider-
ably depending on age (Dooley 1991). Sidewards lighting
using fluorescent lamps or diffusive optical fibres (DOF) for
illuminating cultures produced similar potato plantlets of cv
Benimaru (Kozai et al. 1992). Using DOF reduced cooling
costs, increased PPFD when culture vessels were placed
355
close to the light source, increased space efficiency with the
use of stacked vessels, and increased plantlet leaf area (Kozai
et al.1992). No reports using high-pressure sodium or metal
halide lamps for potato in vitro have been located by the
author of this review.
New Lighting Technologies
Lighting systems that allow for reduced power demand
through the use of hybrid solar and artificial lighting (HYSAL),
and the use of specialized electric lighting systems are in
development (Cuello 2002). Two experimental HYSAL systems
are being tested; one uses xenon-metal halide lamps and the
second uses LEDs. In both experimental systems, light mirrors
and fibre optics are used as the solar conection systems.
Potato is one of the crops being considered for life support in
space (Cuello 2002), and efficiencies in light delivery for in
v/tro systems for potato will undoubtedly result from this
research (Wheeler 2002; Wilson et al. 1993).
Light-emitting diodes (LEDs) are small lamps that convert
electrical energy into light as a form of luminescence. They
can now be used as a radiation source for plants (Bula et al.
1991; Jao and Fang 2004a) and as light sources for growth
chambers (Okamoto et al. 1996). Using LEDs increased in
vitro shoot length in response to increasing FR/PPFD ratio
when the R/PPFD ratio was 0.1-0.5 (Miyashita et al.1994, 1995,
1997). Early LEDs had the disadvantage of low emission effi-
ciency (Green et al. 2001).
Pulsed or intermittent light has been considered for illu-
minating plants (Jao and Fang 2004b; Sager and Giger 1980). A
German company, Oellerich GmbH Berlin, recently developed an
intermittent light technology called Chopper-Light-Technology,
which provides considerable s a ~ g s in energy costs with no
reduction in the productivity of shoot cultures of several
woody species (Pinker 2002). Light cycles other than photope-
riodic cycles have been tested on potato plantlets in vitro
(Hayashi et al. 1994). The authors tested 16 h light/8 h dark, 4
h light/1 h dark, 1 h light/0.5 h dark and 0.25 h light/0.25 dark
such that the ratio of light to dark period was 2:1 in all treat-
ments. The amount of electricity consumed for illumination
per 24-h period was the same for all treatments. Dry weight
increase of the plantlets was greater in the shorter light cycle
treatments. Light-emitting diodes at 180 Hz (5.5 ms) and 59%
duty ratio and 16-h light period were recommended for opti-
mal growth of potato plantlets (Jao and Fang 2004a). Natural
light used to illuminate potato tissue cultures in India and
356 AMERICAN JOURNAL OF POTATO RESEARCH
Cuba produced acceptable cultures and considerable savings
of energy (Alix et al. 2001; Perez-Ponce et al. 2000).
Norton et al. (1988) applied a light pipe in conjunction
with metal halide or high pressure sodium lamps to illuminate
several ornamental species in vit)~o and reported that shoot
length was affected. Kozai et al. (1992) used a prototype diffu-
sive DOF light sources in which only photosynthetically active
radiation (PAR) is emitted to illuminate in vitro Benimaru
potato plantlets. Plantlets had shorter stems and increased dry
weight, leaf area, and net photosynthetic rate, all improved
characteristics for micropropagated plants. New high-intensity
lamps powered by microwaves have improved spectral com-
position for plant growth, but reportedly have yet to be used in
controlled environment chambers (Krizek et al. 1993).
Delivery o f the Light Source
Reflective surfaces of light ~ e s as well as incident
light on the culture shelf can be factors influencing growth of
cultures. Most laboratories illuminate plant tissue cultures in a
horizontal plane from above the cultures. However, significant
differences in the fresh and dry weight, leaf area, and shoot
length of potato cv Benimaru were reported when stacked ctfl-
ture vessels were illuminated using fluorescent lamps with a
sideward lighting system (Hayashi et al. 1992; Kitaya et al.
1995; Kozai et al. 1991).
Light a n d Media Constituents
Photochemical changes caused by light degraded
(unchelated) iron (Fe) and caused the oxidation of EDTA in
culture media (Albano and Miller 2001; Hangarter and
Stasinopoulos 1991; Stasinopoulos and Hangarter 1990). Light
degradation of growth regulator components of plant tissue
culture media has been reported for indole acetic acid (Dunlap
and Robacker 1988; Nissen and Sutter 1990; Posthnmus 1971;
Yamakawa et al. 1979) and abscisic acid (Smith 1992). A com-
bination of light and Murashige and Skoog (1962) salts at pH of
5 was partially alleviated by raising the pH of the media to 7.0
(Dunlap and Robacker 1988). Light can react with the antibi-
otic chloramphenicol to produce breakdown products delete-
rious to plant tissues in culture (Barg et al. 1983a, 1983b). Care
should therefore be taken when storing plant tissue culture
media, and when anomalous results are observed deteriora-
tion of media coltstituents should be considered.
Vol. 82
MODIFYING GROWTH A N D
D E V E L O P M E N T OF POTATO
I N V I T R O WITH LIGHT
Surprisingly few experiments have reported testing the
effect of irradiance levels and spectral quality of light on mor-
phogenesis of potato in v/tro (Bajaj and McAllan 1969; Sarkar
et al. 1996; Seabrook 1987; Seabrook and Douglass 1998; Wilson
et al. 1993). Many authors simply state the environmental con-
ditions employed with no mention of the testing of any other
levels. In view of the reported photomorphogenetic control of
shoot regeneration in vitro in tomato (Lercari et al. 1999), and
grape (Ch~e and Pool 1989), and control of axillary shoot
growth of tomato in vivo (Tucker 1975), it is surprising that lit-
fie work on the effects of light on shoot growth and regenera-
tion in potato has been reported. Light strongly promotes gene
transfer from Agrobacterium to plant cells of Arabidopsis
thaliana and Phaseolus acutifolius in vitro (Zambre et al.
2003). The formation of callus from stem internode explants of
potato and cell suspensions required a reduction in irrradiance
from 600-1,500 lux to 500-600 lux (Cassells and Long 1982). A
higher irradiance reduced callus colony formation by 80%.
Irradiance
Irradiance, or photon flux density available to plant tissue
cultures, is controlled by the light source, the distance of the
cultures from the source, and the shape and type of material
used to fabricate the culture vessel (Fujiwara and Kozai 1995).
Shading of the cultures by closures can be a significant vari-
able (Fujiwara and Kozal 1995), and the type of closure used
should be carefully monitored. Shading of cultures in test
tubes can be partly alleviated by the use of slanted test tube
racks. Fluorescent lamps provide a reasonably even horizontal
illumination if they are placed some distance (15-20 cm) from
the cultures. A light meter should be used to test for uneven
photon flux density across the shelf and the position of the
lamps adjusted accordingly.
Novgtk and Zadina (1987) used 10,000 lux to produce
plants with "compact shoot growth and an expansion of the
leaf blade area." Struik and Wiersema (1999) cultured plantlets
under 800 to 500 lux. Thus, it appears that increasing leaf area,
perhaps through the use of high irradiance levels or various
spectral qualities, should be investigated as a means of manip-
ulating growth and mo~]ohogenesis. End-of-day light treat-
2005 SEABROOK: LIGHT AND POTATO I N VITRO
ments with red and far-red light reduced transplant height and
total leaf length of tomato plants grown in controlled environ-
ments (Decotean and Friend 1991). However, no similar work
on potatoes has been located, and it would be interesting to
determine ff red and far-red light applied to i n vitro plantlet
cultures had a similar effect. High light intensity (120 pmol- m -2
/s -1) with a 12-b photoperiod was the most efficacious treat-
ment for the rooting of in vitro-grown cuttings of potato
(Wang and Hu 1985). Generally, in a medium lacking growth
regulator, roots form readily on potato plantlets in vitro, and
no other measures are necessary.
The capacity of photosynthetic tissues of potato to tolerate
high light irradiances (375 ~mol' m-2s-l) can be mediated by low-
ering the temperature at which in vivo plants are grown (Stef-
fen and Palta 1989). Perhaps acclimatization of tissue cultured
plantlets to conditions of high irradiance can be improved by
lowering the temperature in vitro for similar time periods.
Light Period and Vegetative Growth
A 16-h photoperiod is necessary to maintain vegetative
growth of potato plantlets i n vitro (Dodds et al. 1992),
although this is possibly an effect of a high DLI rather than
photoperiod. Potato plantlet cultures are usually maintained in
a 16-h photoperiod and an irradiance of 6,000 lux (Heszky and
Nagy 1987). A 16-h photoperiod and an irradiance of ca. 1,000
lux (or 90 ~E m -z s -l) (sic)(lAndsay 1987); 2,000-3,000 lux (sic)
(Tao et al. 1987) are generally recommended for the growth
and morphogenesis of meristem tips (Lindsay 1987; Nov~ik and
Zadina 1987; Wang and Hu 1985). Initiating material from etio-
lated tuber sprouts for plantlet cultures is common, and gen-
erally a 14~h photoperiod with an irradiance of 150 ~tE s-lm-2
(sic) is used (Chen and Li 1987; Lindsay 1987).
Hussey and Stacey (1981) reported increased node for-
mation in multiplication cultures with continuous light. How-
ever, as a potato extract was used in the medium, this might
have influenced the results. Tissue-cultured potato plantlets
grown in controlled environment chambers under 24-h illumi-
nation exhibit chlorosis and necrotic spotting (Cao and Tib-
bitts 1991; Cushman and Tibbitts 1996). In the author=s
laboratory potato plantlets grown in vitro under continuous
light do not thrive.
Photoautotrophic Growth In Vitro
Photoautotrophic growth i n vitro is the application of
very high (=150-375 p_mol-2s-1) light irradiance and supplemen-
357
tal CO2 to cultures grown in a medium with little or no sucrose
to supply organic carbon (Pruski et al. 2002; Zobayed et al.
1999). Potato cultures in photoautotrophic conditions exhib-
ited an increase in the number of nodes, dry weight, twice as
many stomata, increased epicuticular wax, and thicker leaves
compared to cultures grown with a lower PPE 20 g/L sucrose
in the medium, and no supplemental CO2 (Zobayed et al. 1999).
An inverse relationship between sucrose concentration and
photosynthetic rate has been reported (Wolf et al. 1998). High
net carbon assimilation rates cannot be sustained in potato
plants grown with a nutrient film technique (Stutte et al. 1996).
The nutrient film protocols used by Stutte et al. (1996)
employed high CO2 levels, long (12 h) photoperiods, and high
light irradiance. Stressed in vitro-grown potato plantlets
exposed to high light and high C Q levels exhibited increased
DNA methylation (Joyce and Cassells 2002). It is possible that
increased DNA methylation is linked to the stress of high net
carbon assimilation rates exhibited by in vitro-grown
plantlets. The saturation irradiance for field-grown potatoes is
1,200 ~tE/m2/s (sic), which is ca. 60% of full sunlight (Dwelle
1985), and researchers culturing potatoes in vitro in photoau-
totrophic growth conditions could use this figure as a guide to
the maximum irradiance necessary for potato under these con-
ditions.
Spectral Quality
Spectral quality of light is the relative intensity and quan-
tity of the different wavelengths emitted by a light source and
perceived by photoreceptors within the plant. Cool-white or
Grohix fluorescent lamps are commonly used for potato tissue
culture (Lindsay 1987; Schilde-Rentschler and Schmiediche
1984; Tao et al. 1987). Lamps vary in spectral quality (Figure 1)
(Anderson 1986; Pennazio and Redolfi 1973; Seabrook 1987;
Wilson et al. 1993), and morphogenesis and growth can be
affected (Seabrook 1987; Wilson et al. 1993). Increasing the red
portion of the spectrum enlarged leaf area of tissue-cultured
potato plantlets (Charles et al. 1992; Garner and Blake 1989;
Seabrook 1987). Significant differences in number of leaves,
leaf area, stem length, and (lit weight of potato cvs Shepody
and Caribe grown under cool-white/Grolux lamp combina-
tions were recorded by Seabrook (1987), Kadkade and Wether-
bee (1983) and Marks and Simpson (1999). Axillary branching
in potato plantlets in vitro in response to different light
sources, e.g., Grolux fluorescent and cool-white + Grohix flu-
orescent (Seabrook 1987) is probably a response to R and FR
358 AMERICAN JOURNAL OF POTATO RESEARCH Vol. 82

light (Heins and v~rflkins 1997; Tucker 1976). Fluorescent lamps
with balanced spectral wavelengths accounting for the need
for photosynthetic photon flux efficacy and R/FR photon flux
ratio are needed for plant tissue culture (Murakami et al.
1992). Short pulses of red light applied to potato sprouts
shortly after excision from the tuber promoted tuberization in
vitro, and treatment with far-red light delayed tuberization
(Blanc et al. 1986), possibly indicating that the active form of
phytochrome was present in the tubers.
Management of plant growth in greenhouse crops using
spectral filters has been reported (Decoteau et al. 1993;
Mortensen and Stromme 1987), and only recently have spec-
tral filters been used in vitro. Photo-selective films have been
used to control the light spectrum and modify plant growth in
greenhouses and controlled environments (Kuboda et al. 2000;
Tsekleev et al. 1992; Tanaka et al. 1992; Van Haeringen et al.
I ~
0 2- ( ~
1
l ;f t
01- ~ I
0
1998). Orchid plantlets cultured in a vessel made of 25-Fro
Neoflon PFA film were larger and more vigorous than plantlets
grown in test tubes (Tanaka et al. 1992). Seabrook and Dou-
glass (1998) used yellow plexiglass filters, which removed blue
light (380 to 525 nm) (Figure 2), to modify the growth of potato
plantlets in tissue culture. Eliminating blue light portions of
the spectrum increased plantlet height (Figure 3) and reduced
the incidence of oedemas (intumescences), small pale yellow
protuberances on leaves and stems which can affect large
areas of the cultured plantlet (Seabrook and Douglass 1998).
Specific wavelengths of light have therefore been implicated in
the disruption of water relations within the plant tissues, lead-
ing to the formation of intumescences in vivo (Rangarajan and
Tibbitts 1994) and in vitro (Seabrook and Douglass 1998).
Plant Growth Regulators a n d Light In Vitro
Interesting synergistic effects between IAA and R light on
in vitro single-node cuttings of potato cv Miranda were
AGRO
...... GROULX reported by Aksenova et al. (1994). In the presence of R light,
IAA influenced stem length reduction and the induction of
tuberization. Further, Aksenova et al. (1994) noted that kinetin
in the presence of blue (B) light strongly promoted tuber for-

0.2 - ~
i t - CWF * AGRO
~ i / "-~ ...... CWF . GROULX
Spectral Comparison Between Two Filters

60.0
o

0.6 50.0
INCAN Filters
E ..... TL-84
0.2
40.0
== ....
- ;"2
,~ o.1 E
¢- c~ 30.0
._m E
0 f, i , i , , , ,
iI i, /%
0.2- it
,' ,,
* •
• \
•
: J "-,, CWF * INCAN

_A
: : " - ..... cw~
10.0
I s

° %; . . . . . ';;O-
I I I = I
WAVELENGTH - - - nm
4oo flOO 800 1ooo
nm
FIGURE 1.
Spectral output ( n m ) of five common lamps used to illuminate Wavelength
potato cultures. AGRO = Agrolite fluorescent, GROLUX =
Glolux fluorescent, CWF = cool-white fluorescent, INCAN = FIGURE 2.
60w incandescent lamp, TL-84 = Philips TL 40w/84RS fluores- Comparison o f spectral wavelength o f light from cool-white flu-
cent lamp. Differences in growth o f plantlets illuminated by orescent lamps filtered through clear and yellow plexiglass ill-
various lamps were recorded (Seabrook 1987). ters (Seabrook and Douglass 1998).
2005 SEABROOK: LIGHT AND POTATO I N VITRO
mation and increased total fresh weight as well as increasing
the root + stolon: haulm ratio, apparently through the conju-
gation of IAA with glucose (Aksenova et al. 1990). Auxins and
cytokinins apparently mediated the morphogenetic effects of
light on potato in vitro (Sergeeva et al. t994).
Red light has been implicated in the synthesis of chloro-
phylls; however, Sarkar et al. (1996) discerned differences in
the chlorophyll content of two cvs (Kufri Jyoti and Kufri Chan-
dramukhi) illuminated with fluorescent and incandescent
lamps fitted with filters to provide white light enriched with R
and B light. Sarkar et al. (1996) added gibbereUin to the culture
medium, and this may have influenced the results. Red light is
known to prevent leaf abscission in plants, but sub-optimal
levels of R light have been implicated in dark-induced leaf
abscission in mung bean (Decoteau and Craker 1984). Possi-
bly, the use of R light could enhance the retention of leaves in
tissue-cultured potato plantlets. Short pulses (5 rain) of R light
have been implicated in shortening the time to tuberization for
in vitro grown potato sprouts (Blanc et al. 1986), and R light
activated the elongation of potato stems (Konstantinova et al.
1987). Concurrent blue and red light provided at the beginning
of the photoperiod by LEDs resulted in an increase in fresh/dry
Yellow Filter Effects on Potato Plants
~ntrOI
w filter
10
E
f.-
c 5
m Blue light increased the number of axillary buds, and its
Brador Russet Burbank Shepody
FIGURE 3.
Effect of yellow filter on plantlet length o f three potato culti-
v a r s g r o w n i n v i t r o . Leaf area from both treatments remained
the same (Seabrook and Douglass 1998).
359
weight accumulation of potato plantlets in culture (Jao and
Fang 2004b).
Far-red light stimulated the conversion of 1-aminocyclo-
propane-l-carboxylic acid to ethylene in Begonia X hiemalis
leaf and flower discs in vivo (Rudnicld et al. 1993). It would be
interesting to test potato tissues for ethylene production when
illuminated by various monochromatic light spectra, espe-
cially in view of the sensitivity of potato in vitro to restricted
gaseous exchange (Jackson et al. 1991) and the formation of
oedemas under certain light regimes (Rangarajan and Tibbitts
1994; Seabrook and Douglass 1998). The motivation for the
Seabrook and Douglass (1998) study was the unacceptably
short, rosette-type growth habits of certain potato cutivars
such as AC Brador and Shepody so that multiplication rates
were adversely affected. The use of a yellow plexiglass or cel-
lophane filter increased stem length by 200°/5while maintaining
the number of leaves and increasing leaf area. An additional
benefit of removing blue light was the reduction in the inci-
dence of oedemas (Seabrook and Douglass 1998). Aksenova et
al. (1994) noted that B light in the presence of kinetin strongly
stimulated potato tuber formation, increased fresh weight, and
increased root + stolon to shoot ratio. Challakhyan et al.
(1992), working with S. andigenum (sic) i n vivo, compared
growth and morphogenesis under both B and R light and
reported that ilhunination with blue light increased leaf area,
"total mass (g)," and photosynthesis.
Irradiation with UV light induced the biosynthesis of the
stilbene phytoalexin trans-resveratrol in the leaves of both i n
vitro and greenhouse-grown potato plants (Lippmann et al.
2000). Trans-resveratrol (3,5,4'- trihydroxy stilbene) is reported
to have anti-fimgal qualifies, and could be of use against fimgal
pathogens of potato (Lippmann et al. 2000). It would be useful
to know if trans-resveratrol could be used to partially protect
potato from tissue culture plantlets from pathogens during
acclimatization to the field.
influence depended on the fluence rate whereas R light influ-
ences were independent of fluence rate and reduced correla-
tive inhibition (apical dominance) (Muleo et al. 2001). Light
spectral quality could be used to manipulate both leafy shoot
and plantlet production in potato rapid multiplication regimes.
Altering the spectral wavelength of lamps could change cor-
relative inhibition (apical dominance) of the plantlet to pro-
duce more than one microtuber per whole plantlet.
360 AMERICAN JOURNAL OF POTATO RESEARCH
Microtubers exposed to light during induction and early
development tend to have well-developed periderm, are fre-
quently green, and have improved disease and dessication
resistance (Naik and Sarkar 1997). When microtubers were
produced in light, more eyes were formed, survival was
improved, and growth in the field of the subsequent plantlets
was more vigorous (Gopal et al. 1998). Martlnez-Garcia at al.
(2002) used S o l a n u m t u b e r o s u m ssp. a n d i g e n a , an obligate
SD plant, to determine the interacting role of photoperiod and
gibberellins in tuberization. They propose that tuberization in
potato is similar to the flowering process in A r a b i d o p s i s : t w o
separate pathways control tuber formation--a photoperiodic-
dependent and a gibberellin-dependent pathway. The induc-
tion of microtuberization i n v i t r o can be used to assess the
maturity group (i.e., cultivar reaction to photoperiod) of
potato germplasm (Lentini and Earle 1991; Pelacho et al. 1994;
Veramendi et al. 2000).
The spectral quality of light transmitted through various
culture vessels can vary (Dooley 1991). Polycarbonate did not
transmit light wavelengths less than 390 rim, glass <290 nm,
and polystyrene <300 nm (Dooley 1991). Researchers should
therefore be aware that some morphogenetically active wave-
lengths of light could be excluded from the illumination pro-
vided to cultures. The gaseous content in culture vessels
containing potato plantlets was not affected by changing the
light conditions (Buddendorf-Joosten and Woltering 1996).
FIGURE 4.
Examples of microtu-
bers formed on in vitro
a b
single-node cuttings in
r e s p o n s e to various
photoperiodic treat-
2 ments: ( a ) desirable
microtuber for multi-
pliation; ( b ) microtu-
ber subtended
stolon; ( c ) secondary
by
microtuber formation;
(d) secondary axillary
microtubers. Leave
were removed from the
cuttings so that draw-
ings could be prepared
I I
1¢m ( S e a b r o o k et al. 1993).
Vol. 82
Photoperiod
Tuberization of potato can be promoted by environmental
(photoperiod) and by developmental (growth regulators) fac-
tors (Martlnez-Garcia et al. 2002). This may explain the appar-
ently contradictory information in the literature regarding the
effect of photoperiod on tuberization i n v i t r o noted by Ewing
and Struik (1992). Early studies on tuberization in v i t r o used
etiolated sprout sections from field-grown tubers in a culture
medium containing high sucrose levels (Barker 1953;
Lawrence and Barker 1963). Later reports studying photoperi-
odic effects on microtuberization also used etiolated sprout
sections (Pelacho and Mingo-Castel 1991). Microtuberization
of etiolated sprout sections is independent of light (Lawrence
and Barker 1963; Pelacho and Mingo-Castel 1991). The use of
single-node cuttings excised from tissue-cultured plantlets is
more common and avoids the influence of the tuber tissue
from which sprout sections originate (Hussey and Stacey 1981;
Leclerc et at. 1994; Levy et aL 1993; Seabrook et al. 1993). Dif-
ferences in the physiological state of stock material may
account for some of the varying results reported.
The photoperiodic reaction of the potato plant is well-
known (Chailakhyan et al. 1982; (Chailakhyan et al. 1992;
Driver and Hawkes 1943; Ewing 1990; Mendoza and Haynes
1977). Photoperiod is a major morphogenetic control of tuber-
ization in potato in tissue culture (Coleman and Coleman 2000;
Seabrook et al. 1993). Potato plantlets i n v i t r o can be induced
to produce microtubers by inductive short daylengths (Garner
and Blake 1989; Lentini and Earle 1991; Pelacho and Mingo-
Castel 1991; Seabrook et al. 1993). Tuberization i n v i t r o can be
induced in the absence of high sucrose levels, with no growth
regulators and using inductive daylengths (Garner and Blake
1989). Ewing and Wareing (1978) reported that the amount and
type of tuberization on cuttings i n vivo was determined by the
amount and type of tuberization stimulus, and Seabrook et al.
(1993) demonstrated that the morphology (Figure 4) of tuber
formation i n v i t r o was influenced by photoperiod. Time to
tuberize (Veramendi et al. 2000), number of microtubers
(Lentini and Earle 1991; Pelacho et al. 1994) and Adegree of
tuberization@ (Veramendi et al. 2000) were used to categorize
potato cultivars into maturity groups.
Photoperiodic control of tuberization in potato tissues cul-
tured i n vitro has been established (Dobrfinski 2000; Garner
and Blake 1989; Hussey and Stacey 1984; Lentini and Earle
1991; Martlnez-Garcfa et al. 2002; Seabrook et al. 1993). The
use of media supplemented with gibberellins in the report by
2005 SEABROOK: LIGHT AND POTATO I N VITRO 361

Martinez-Garcia et al. (2002) may confound the results of

experiments with regard to light requirements. Struik and
Wiersema (1999) recommend incubation in the dark or at low
W
light intensities (100-50 hix) with a light period of 8 h to induce
microtubers. Lindsay (1987) reported incubating cultures to m
produce microtubers at a 16-h photoperiod and diffuse light
(10 ~E m-2s-1). However, low irradiance levels can delay in
vitro tuberization (Jackson 1999). As a general nile, microtu-
berization can be initiated by incubating cultures in darkness 0
or by exposure to a 10. to 12-h photoperiod. 100
The involvement of phytochrome in tuberization in vivo 8O
has been established (Batutis and Ewing 1982; Yanofsky et al.
2000). When environmental signals such as photoperiod are
m
used to promote microtuberization (Blanc et al.1986; Jackson
1999) phytochrome may be involved. Blue light (4000-580 nm)
promoted more tuberization than red light (600-700 nm) when
applied for 16 h photoperiods (Aksenova et al. 1989). 0

Microtuberization
8O
Exposure of potato plantlets from single-node cuttings to
8 h (SD) and 16 h (LD), and then subsequent single-node cut-
tings to SD and LD indicated that the production of microtu-
4o
bers was strongly influenced by genotype and photoperiod.
Incomplete photoperiodic induction of microtubers resulted 20
in secondary tubers, the production of stolons, and the forma-
0
tion of additional tubers in the axils of the leaf of single-node
100
cuttings (Figure 4) (Seabrook et al. 1993). Potato cultivars
(Jemseg, Kathadin, Russet Burbank, and Superior) varied in 80
reaction to photoperiodic treatments with short days (8 h
light) generally being the optimum treatment (Figure 5) •)i
==-)+
(Seabrook et al. 1993).
The effects of light on the greening of field-grown potato
2O
tubers are well known (Baerug 1962). However, Nalk and
Sarkar (1997) reported that light-induced greening of microtu-
1 2 3 4
bers can be used to improve the storage of microtubers, which LD-LD LD-$D SD-LD $D-$D
otherwise tend to loose weight and shrink. A 16-h photoperiod Photoperiodic treatment
at an h-radiance level of 30 ~tmol m-2s-1 for 10 days improved
miorotuber type
biomass loss during storage, and subsequent sprout emer-
gence (Naik and Sarkar 1997). m. mo +° Do
Light irradiance affects the duration of dormancy of F I G U R E 5.
potato microtubers (Tgtbori et al. 2000); the lower the irradi- P e r c e n t a g e o f various t y p e s o f m i c r o t u b e r s ( s e e Figure 4) pro-
duced by p o t a t o single-node cuttings in vitro e x p o s e d to pho-
ance, the longer the dormancy of cvs Desir6e and Gtilbaba. toperiodic treatments. Plantlets e x p o s e d to LD = 16 h light; SD =
Microtubers induced and developed in an 8-h photoperiod 8 h light. Single-node cuttings from p l a n t l e t s e x p o s e d to LD a n d
SD and g r o w n in tuberization medium. Microtuber types: a = nor-
compared with darkness exhibited shorter dormancy (Cole- r0~l; b = microtuber s u b t e n d e d by stolon; c -- s e c o n d a r y microtu-
man and Coleman 2000; Gopal et al. 1998). Microtubers stored ber; d = secondary axilhry m i c r o t u b e r ( S e a b r o o k e t al. 1993).
362 AMERICAN JOURNAL O F POTATO RESEARCH
under diffuse light had longer dormancy than microtubers
maintained in the dark (Gopal et al. 2004).
Virus E r a d i c a t i o n f r o m Shoot Tips
Subsequent to excision of the meristem including several
leaf primordia, dark incubation is not advised (Huth and Bode
1970). Shoot explants excised from potato sprouts on tubers
maintained in light (5.0 W m -2) grew better than those grown
in darkness (Casseis and Long 1982). A photoperiod of 16 h
rather than 8 h is recommended (Cassells and Long 1982; Mel-
lor and Stace-Smith 1987; Nov~k and Zandina 1987). Although
low light intensity (irradiance) (100 lux) is necessary to pre-
vent browning of the tissues when incubating newly excised
shoot tips, after 4 wk of incubation, light intensity can be
increased considerably to 2 to 4 kilolux (Mellor and Stace-
Smith 1987; Wang and Hu 1985). Very high light intensities (30
kilolux) have been reported to be useful for incubating shoot
tips (Klein and Livingston 1983). All recommendations of irra-
diance levels have to be noted with the caveat that both the
light source (lamp) and the method of measurement have to be
carefully noted to elicit any meaningful comparisons (Hart
1988). Pemmzio and Redolfi (1973) reported the effect of dif-
ferent light spectra on the growth and development of shoot
tips of potato.
In Vitro Conservation
Potato germplasm is maintained at 1000 lux at a 16-h pho-
toperiod at 6 C at the International Potato Centre (C.I.P.) in
Lima, Peru (Dodds et al. 1991). However, Westcott et al. (1977)
recommended a 16-h photoperiod, a light intensity of 4,000 lux
at 22 C with monthly transfers to fresh medium for storage of
potato germplasm, or the same conditions with storage at 6 C
and less frequent transfers of tissues. Heszky and Nagy (1987)
maintained single-node cuttings in vitro at 6,000 lux under a
16-h photoperiod. Light affected the viability of potato cultures
maintained at low temperatures (Pruski et al. 2000). At the
Canadian Potato Gene Repository at the Potato Research Cen-
tre, Fredericton, New Brunswick, plantlet cultures are main-
tained under a 16-h photoperiod with an irradiance of 80
~unol'm-2s-~ combined with lower temperatures (12 C) for
selected cultivars. It is surprising that light irradiance levels
and spectral quality do not seem to have been investigated as
a means of control of growth and development in germplasm
conservation regimes.
Vol. 82
Pre-treatment o f P o t a t o Tissues In Vitro
Although light treatments can be used to improve the effi-
ciency of in vitro propagation regimes (Economou and Read
1987), there are few reports on pre-treatment of potato tissue
with light. Seabrook et al. (1993) reported that long day expo-
sure of plantlets prior to tuber induction conditions and short
days during tuberization increased yield and quality of micro-
tubers in cvs Katahdin and Russet Burbank, but had little
effect on cvs Jemseg and Superior. Pre-treatment of tissue cul-
ture plantlets in vitro with short photoperiods (8 or 12 h light)
prior to planting them in greenhouses for the production of
minitubers increased yield (Seabrook et al. 1993; Seabrook et
al. 1995).
A c c l i m a t i z a t i o n o f In Vitro Plantlets to
E x Vitro Conditions
High light conditions (1,000 to 3,000 lux) have been rec-
ommended for preparing tissue culture plantlet for acclima-
tion to soil (Murashige 1977). Kozai et al. (1988) tested three
different irradiance (PPF) levels and determined that potato
plantlets at the highest level (400 l~mol m-2s-1) were most vig-
orons and developed sufficient roots for acclimatization.
Muraldshi and Harris (1987) also used high irradiance (4,000
lux) levels prior to planting out in the greenhouse. Although
enriching the gaseous atmosphere of cultures may improve
water-use efficiency and increase net carbon uptake, the pos-
sibility of poor root development due to the effects of elevated
CO2 on photosynthesis over a long period of time must be con-
sidered (Chaves 1994). Thus, high irradiance levels and ele-
vated CO2 may have the undesirable side effect of producing a
plant unsuited to the stress of acclimatization. Steffen and
Palm (1989) reported that one way to overcome this high irra-
diance stress exhibited by potato plants in vivo was to grow
them at 12 C for 4 wk. The efficacy of low temperatures prior
to acclimatization for alleviating high irradiance stress of in
vitro potato plantlets should be tested.
CONCLUSIONS
New technologies are required to provide high light irra-
diances for autonomous growth of potato cultures without the
expense of cooling. Lighting systems such as hybrid solar and
artificial lighting (HYSAL), light-emitting diodes (LEDs),
pulsed light (Chopper light), and the use of natural light have
the potential to conserve electrical power. Further research on
2005 SEABROOK: LIGHT AND POTATO I N V I T R O
t h e effects o f t h e interaction o f p h o t o p e r i o d with e x o g e n o u s
a n d e n d o g e n o u s g r o w t h regulators o n microtuberization are
n e e d e d to fully u n d e r s t a n d the tuberization p r o c e s s in potato.
Studies o n the effects o f light o n d o r m a n c y o f m i c r o t u b e r s
w o u l d b e useful for t h e s h i p m e n t a n d storage o f p o t a t o
germplasm. Testing the daily light integral (DQLI) a n d pho-
t o p e r i o d could d e t e r m i n e t h e physiological p a r a m e t e r s under-
lying the r e a c t i o n o f p o t a t o t i s s u e s to light i n vitro. The
w e l l - k n o w n effect o f s e e d s o u r c e environment, climate, and
soil c o n d i t i o n s o n the s u b s e q u e n t yield a n d g r o w t h o f the
p o t a t o crop, leads to the c o n c l u s i o n t h a t the effect o f pre-treat-
m e n t with light (photoperiod, s p e c t r a l quality, a n d irradiance)
o n various a s p e c t s o f p o t a t o tissue culture s h o u l d b e consid-
ered. Very little r e s e a r c h h a s b e e n r e p o r t e d o n the effects o f
specific w a v e l e n g t h s and c o m b i n a t i o n s o f w a v e l e n g t h s o f light
o n productivity o f rapid multiplication cultures, m i c r o t u b e r -
ization and r e g e n e r a t i o n (specifically s o m a t i c e m b r y o g e n e s i s )
o f p o t a t o in vitro.
The relationship b e t w e e n t h e putative effects o f p h o t o p e -
riod a n d DQLI o n p o t a t o t i s s u e s i n vitro s h o u l d b e investi-
gated particularly in view o f the r e p o r t e d effects o f light p e r i o d
on vegetative growth.
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