The  

effect  of  light  quality  on  growth  of  lettuce  

 
Yuta  SAITO*,  Hiroshi  SHIMIZU**  
 Hiroshi  NAKASHIMA***,  Juro  MIYASAKA****,  Katsuaki  OHDOI*****  

*  Graduate  student,  Graduate  School  of  Agriculture,  Kyoto  University,  Sakyo-­ku,  Kyoto  606-­8502,  Japan  
(  e-­mail:  saito@a06nogaku.mbox.media.kyoto-­u.ac.jp,  Tel:81-­075-­753-­6165)    
**  Professor,  Graduate  School  of  Agriculture,  Kyoto  University,  Sakyo-­ku,  Kyoto  606-­8502,  Japan  
(e-­mail:  hshimizu@kais.kyoto-­u.ac.jp.)    
***  Associate  Professor,  Graduate  School  of  Agriculture,  Kyoto  University,  Sakyo-­ku,  Kyoto  606-­8502,  Japan  
(e-­mail:  hiron@kais.kyoto-­u.ac.jp)  
****  Assistant  Professor,  Graduate  School  of  Agriculture,  Kyoto  University,  Sakyo-­ku,  Kyoto  606-­8502,  Japan  
(e-­mail:  miyasaka@kais.kyoto-­u.ac.jp)  
*****  Assistant  Professor,  Graduate  School  of  Agriculture,  Kyoto  University,  Sakyo-­ku,  Kyoto  606-­8502,  Japan  
 (e-­mail:  ohdoi@kais.kyoto-­u.ac.jp)  

Abstract:   Plant   factory   is   a   plant   production   system   with   controlled   temperature,   carbon   dioxide  
concentration  and  light  environment  to  enhance  plant  growth  efficiently.   Fluorescent   lamp  is  popularly  
used  in  a  plant  factory,  however,  the  spectrum  of  fluorescent  light  is  broad  and  some  portion  of  them  is  
not  necessary  for  photosynthesis.  On  the  other   hand,  LED  (light  emitting  diode)  has  narrow  band  width  
and  is  getting   easy  to  use  these   days.  The  final  goal  of  this  project  is   to   develop  a  new  light   apparatus  
with  LEDs  for  plant  factory.  The  specific  objective  is  to  investigate  the  effective  light  quality  for  growing  
plant  more  rapidly.  The  effect  of  light  generated  by  monochromic  and  mixed  radiation  from  LEDs  (blue,  
green,  red)  on  the  response  of  lettuce  (Lactuca  sativa  L.)  were  evaluated  and  compared  with  the  effect  of  
lettuce  irradiated  by  fluorescent  lamps.  Photosynthesis  rate  of  lettuce  increased  in  monochromic  red  light  
and   the   light   mixed   with   red   and   blue   light.   Number   of   leaves   decreased   in   monochromic   blue   light.  
Fresh   weight  of  lettuce   was  increased  in   monochromic  red  and   fluorescent  light,  but   monochromic  red  
light   significantly   decreased   dry   weight   rate.   In   the   view   of   energy   saving,   LED   is   much   better   than  
fluorescent  lamp.  Especially,  red  LED  is  good.    
Keywords:  plant  factory,  light  quality,  carbon  dioxide  concentration,  light  intensity,  morphogenesis,  fresh  
weight,  LED    

germination   and   photosynthesis   (Flint   and   McAllister   1937;;  
1.  INTRODUCTION  
Balegh   and   Biddulph   1970).   Blue   light   effects   stomatal  
Agriculture   and   food   issues   are   getting   known   widely.   For   opening  and  chloroplast  development  (Takemiya  et  al.  2007;;  
example,   residual   agricultural   chemicals,   poor   harvest   by   Akoyunoglou   and   Anni   1984).   Green   light   slows   down   or  
abnormal   weather   conditions   are   often   heard   all   over   the   stop   the   plant   development,   but   it   brings   positive   effects   in  
world.   Especially  in  Japan,   there   are  other  problems  such  as   special  conditions  (Folta  2004,  2007;;  Terashima  et  al.  2009).  
the   increase   of   imported   foods   and   the   decrease   of   farmers.   In   many   plants,   it   has   been   studied   that   the   absorbance   of  
Plant   factory   is   expected   to   be   one   of   the   ways   to   solve   the   leaves   and   the   quantum   yield   of   photosynthesis   concerning  
issues.   The   plant   factory   is   controlled   inside   environment   various   wavelength.   We   may   produce   an   optimum   light  
such   as   lighting,   temperature,   carbon   dioxide   concentration   quality   for   plant   growth   by   combining   the   LED¶s  
and   cleanliness   to   enhance   plant   growth   effectively.   That   is   characteristic  of  light  quality  with  researches  of  plant  growth.  
why  plant  factory  can  produce  a  lot  of  safe  vegetables  in  spite   In   fact,   photosynthesis   rate   and   growth   rate   of   plant   grown  
of  season  or  weather.  However,  the   initial  and  running  costs   under  LED  were  reported  (Mori  and  Takatsuji  2000,  Mori  et  
for   environmental   control   are   very   expensive.   These   al.  2002).  
problems   prevent   a   plant   factory   from   spreading   widely.   In  
 The  final  goal  of  our  study  is   to  improve  lighting  apparatus  
running   cost,   the   cost   of   lighting   for   plant   growth   is   one   of  
for   plant   factories   to   spread   them   widely.   So   the   specific  
the  large  portions.  
objective   of   this   research   is   to   investigate   the   best   light  
Recently  a  light  emitting  diode  (LED)   is  spreading  as  a  new   qualities   for   plant   growth   by   examining   photosynthesis   rate  
light   source.   A   LED   has   good   life   performance,   energy-­ and   growth   response   of   lettuce   (Lactuca   sativa   L.)   radiated  
saving   ability   and   various   and   narrow   bandwidth   of   by  LEDs  and  fluorescent  lamps.  
wavelength.   Combining   several   LEDs   with   different  
bandwidth  enables  to  produce  many  kind  of  light  quality.  The  
effects   of   light   qualities   on   the   growth   and   development   of  
plant   have   been   studied   for   a   long   time.   Red   light   promotes  

         
    
 
h/d  light  period  and  25  °C  in  light  period  and  18   °C  in  dark  
period.  
Table  1.  Light  intensity  rates  of  blue,  green  and  red  LEDs  for  
five  light  qualities  using  LED  systems  (%)  

  Blue  LED   Green  LED   Red  LED  

R   0.0   0.0   100.0  

RB   16.7   0.0   83.3  

B   100.0   0.0   0.0  

SL-­L   33.4   33.9   32.7  

  FL-­L   29.8   39.6   30.7  

Fig.  1.  Developed  LED  system  
2.3    Hydroponics  

Nutrient   solution   pools   of   hydroponics   were   consisted   of  

culture   bed   (730   wide   mm   ×   665   length   mm   ×   170   height  
mm)   covered   tarpaulin   and   culture   panel.   The   quantity   of  
nutrient   solution  in   the  pools  was   maintained  even  by  pump  
and   overflows.   Nutrient   solution   was   kept   pH   5.9±0.2,   EC  
2.00±0.10   by   fertilizer   controller   (CEMCO)   with   acidity  
regulator  (Otsuka)  and  fertilizers  (Otsuka).  
 
2.4    Light  Quality  
Fig.  2.  Spectrums  of  LEDs:  Blue,  Green  and  Red   were  used  
in  this  research.  
 In   photosynthesis   measurement   and   growth   experiment,   six  
2.  MATERIALS  and  METHODS   light  qualities   were   used.  One   was   fluorescent   light  (FL)  set  
by  fluorescent  lamps  and  the  others  were  set  by  LED  systems.  
2.1    Light  Source   The   light   qualities   set   by   LED   systems   were   monochromic  
red   light   (R),   monochromic   blue   light   (B),   the   light   mixed  
We   used   two   light   sources.   One   was   a   fluorescent   lamp   with  red  and  blue  light  (RB),  the  light  imitating  sunlight  (SL-­
(FLR20S・W/M,   TOSHIBA),   the   other   was   a   LED   system.   L),  the  light  imitating  fluorescent  lamp  light  (FL-­L).  Table  1  
The  LED  system  was  consisted  of  blue,  green  and  red  LEDs   shows  light  intensity  rates  of  blue,  green  and  red  LEDs  about  
(LUXEON  Rebel,  Philips  Lumileds)  and  condenser  lenses  as   these  five  light  qualities.  We  referred  to  the  spectrum  data  of  
shown   in   Figure   1.   Each   spectrum   of   LEDs   is   shown   in   sunlight   (Air   Mass   1.5:   ASTM   G173,   NREL)   and   a  
Figure   2.     Photosynthetic   photon   flux   density   (PPFD)   from   fluorescent   lamp   (FLR20S・W/M,   TOSHIBA)   to   set   SL-­L  
LEDs  system  can  be  controlled  by   adjusting  currents   from  a   and  FL-­L.  
DC  power  supply.    This   paragraph   explains   the   reasons   why   we   used   these   six  
light  qualities.  Fluorescent  lamps  are  popularly  used  in  plant  
2.2    Plant  Materials  and  Growth  Condition   factories,  so  FL  was  used  to  reproduce  plant  growth  in  plant  
factories.   R   was   used   because   it   was   reported   that  
Lettuce   (Lactuca   sativa   L.   cv´Greenwave´,   Takii   Seed   Co.,   monochromic   red   light   irradiation   showed   the   highest  
Ltd.)  plants  were  grown  in  urethane  sponge  with  water  under   photosynthesis   rate   in   many   monochromic   light   irradiations  
fluorescent   lamps   for   one   week.   PPF   of   fluorescent   lamps   (Balegh   and   Biddulph   1970;;   Björkman   1973).   Previous  
was   set   at   200   µmolm-­2s-­1.   After   one   week,   lettuce   plants   research  reported  dry  weight  of  plant  grown  increased  in  blue  
were   transplanted   to   hydroponic   system.   We   conducted   two   light  (Hirai  et  al.  2006).  That  is  why  B  was  used.  For  RB,  it  
experiments   namely   photosynthesis   rate   measurement   and   was   reported   that   the   light   mixed   with   red   and   blue   light  
growth   experiment.   In   photosynthesis   rate   measurements,   promoted   plant   growth   and   photosynthesis.   To   reproduce  
transplanted  plants  were  grown  under  fluorescent  lamps  for  a   plant   growth   in   sunlight   which   plants   have   adapted,   SL-­L  
month  (11/26-­12/27,  2009).  In  growth  experiment,  they  were   was   used.   FL-­L   was   used   to   research   the   effect   of   the   light  
grown   under   six   light   qualities   from   fluorescent   lamps   and   which  removing  ultraviolet  and  infrared  rays  from  fluorescent  
LED  systems  for  four  weeks.  All  of  the  plants  were  grown  in   light.  
a  growth  chamber  (KCLP-­1000ⅠCT,  NKsystem)  set  at  a  16  
   

         
    
 
9 treatment   was   lower   than   that   of   RB   treatment   (Mori   and  
Takatsuji   2000).   But   our   result   was   opposite   of   previous  
Photosynthesis  rate  

8
7 result.   It   could   be   due   to   the   difference   plant   specie,  
(µmol  m-­2  s-­1)

6 experiment   environment   and   growth   condition.   The  

5 photosynthesis   rates   under   SL-­L,   FL-­L   and   FL   were   not   so  
4 different.   This   could   be   due   to   little   difference   among   their  
3 light  qualities.  In  spite  of  light  intensity  or  CO2  concentration,  
2 the   result   that   the   highest   photosynthesis   rate   was   in   R,  
1 followed   in   RB   and   in   the   others   in   this   order   was   not  
0 changed.  
R RB B SL-­L FL-­L FL
3.2    Growth  Experiment  
Fig.   3.   Relation   between   light   qualities   and   photosynthesis  
rates:  This  data  was  measured  in  the  case  when  light  intensity  
3.2.1  Number  of  leaves  
was  set  at  200  ρ‘Žିଶ • ିଵ  and  CO2  concentration  was  set  at  
600  ppm.   Figure  4  shows  the  relation  between  light  quality  and  number  
of   leaves.   Large   standard   errors   can   be   seen   in   R   and   FL  
2.5    Photosynthesis  Rate  Measurement   treatments.   Plants   grown   under   R   treatment   had   long   leaves  
and   petioles,   so   that   overlapping   of   leaves   happened.   The  
LI-­6400   (LI-­COR)   was   used   for   photosynthesis   rate   overlapping  makes  difference  to  the  light  quantity  supplying  
measurement.  Plant  leaf  was  fastened  with  a  leaf  chamber  of   plants   and   plant   growth   like   number   of   leaves.   The   light  
LI-­6400  whose  CO2  concentration  set  at  400  ppm  and  applied   intensity  of  the  FL  treatment  is  different  among  transplanted  
one  light  quality  whose  light  intensity  was  set  at  300  µmolm-­ spots   because   the   light   intensity   of   fluorescent   lamp  
2 -­1
s   for   an   hour.   After   confirming   photosynthesis   rate   was   decreases   near   edge   of   the   lamp   tube.   It   could   cause   the  
stable,  the   measurement   was   started.  The  light  intensity   was   difference   of   individual   growth   under   FL.   Other   light  
set  at  300  (250  in  FL)  µmolm-­2s-­1  at  first,  followed  200,  150,   qualities  happened  less  overlapping  of  leaves  than  R  and  had  
100,   80,   60,   40,   20,   10   µmolm-­2s-­1.   In   each   light   intensity,   the   same   light   environment   among   individuals   because   light  
CO2   concentration  was  set  from  0  ppm  to  1500  ppm  per  100   sources  set  right  above.  So   standard  errors  in  their  treatment  
ppm   and   photosynthesis   rate   was   measured   and   stored   at   were  significantly  low.  
every  condition.   Number   of   leaves   was   increased   in   SL-­L   and   significantly  
After   measurement   was   completed   to   10   µmolm-­2s-­1,   light   decreased   in   B.   Plants   grown   in   SL-­L   treatment   had   more  
quality   was   changed   to   the   next   and   the   same   measurement   new   and   small   leaves   than   others.   Decreasing   leaves   of  
procedure  was  repeated.  In  all  measurement,  the  same  area  of   lettuces  grown  under  blue  light  has  been  reported  previously  
the  same  leaf  of  the  same  plant  was  used.   (Hirai   et   al.   2006).   This   research   reported   that   the   effect   of  
blue  light  on  decreasing  leaves  has  the  significant  difference  
2.6    Growth  Experiment   among   monochromic   blue,   blue-­green,   green   and   red   light  
treatment.   And   in   our   research,   this   effect   of   blue   light   has  
the   significant   difference   among   B,   R,   RB,   SL-­L,   FL-­L   and  
This   experiment   had   six   treatments   with   six   light   qualities.  
FL   treatment.   From   these   researches,   it   is   considered   that  
One   treatment   with   nine   plants   was   considered   as   a  
blue  light  could  decrease  number  of  leaves  and  green  and  red  
replication.  But  one  plant  in  R  treatment  and  two  plants  in  FL  
lights  could  suppress  this  decreased  effect  of  blue  light.  
treatment   were   not   grown   well,   so   they   were   removed   in  
replications.   Growth   responses   were   determined   to   measure    
fresh  weight,  dry  weight  and   number  of  leaves  per  plant.  To  
compare  the  costs,  we  measured  electric  power  consumptions.   25.0
The   number   of   leaves   was   measured   all   leaves   contained  
Number of leaves

dead  or  very  small  new  leaves.  The  dry  weight  was  measured   20.0
(leaves/plant)

after  drying  the  samples  in  a  hot  air  oven  at  70  °C  for  4  days.  
15.0
3.  RESULTS  and  DISCUSSIONS  
10.0
3.1    Photosynthesis  Rate  Measurement   5.0

Figure   3   shows   the   relation   between   light   quality   and   0.0

photosynthesis   rate.   Photosynthesis   rate   increased   in   R RB B SL-­L FL-­L FL
monochromic   red   light   (R)   and   RB.   These   light   qualities    
contained  more  red  light  than  others,  so  it  was  considered  that   Fig.  4.  Relation  between  light  quality  and  number  of  leaves:  
red   light   could   promote   photosynthesis.     This   response   has   Each  value  is  the  mean  ±  standard  errors  of  replications.  
been   reported   previously   in   many   species   (Emerson   and  
Lewis   1943;;   Balegh   and   Biddulph   1970;;   Björkman   1973).  
Another   research   reported   that   photosynthesis   rate   of   R  

         
    
 
100 2.5 140

Production efficiency (g/W)

Power consumption (W)

Production efficiency (g/W)
80 120
Fresh weight (g)

2 Power consumption (W)

100
60 1.5 80
40 1 60
40
20 0.5
20
0 0 0
R RB B SL-­L FL-­L FL R B FL
   
Fig.  5.  Relation  between  light  quality  and  fresh  weight:  Each   Fig.   7.   Relation   among   light   quality,   production   efficiency  
value  is  the  mean  ±  standard  errors  of  replications.   and  electric  power  consumption  
3.2.2  Fresh  weight  and  Dry  weight   It   is   considered   R   treatment   could   suppress   dry   matter  
production   for   fresh   weight.   We   think   the   reason   why   dry  
We   measured   fresh   and   dry   weight   of   plant   parts   above   a  
weight   rate   decreased   in   R   could   be   a   bad   plant   posture   of  
culture   panel.   Figure   5   shows   the   relation   between   light  
light  absorption  caused  by  overlapping  of  leaves  and  hanging  
quality   and   fresh   weight.   Large   standard   errors   of   fresh  
down  leaves.  Figure  6  also  shows  that  the  higher  rate  of  red  
weight  can  be  seen  in  R  and  FL  treatments.  It  was  caused  by  
light   a   light   quality   had,   the   less   dry   weight   of   plant   grown  
the   difference   of   competition   or   light   environment   among  
under  the  light  quality  became.  
individuals   as   seen   in   number   of   leaves.   Since   plant  
photosynthesizes  to  grow  up,  it  is  considered  the  light  quality   3.2.3  Comparison  of  electric  power  consumption  
which   shows   high   photosynthesis   rate   produces   heavy   fresh  
Figure   7   shows   the   relation   among   light   quality,   production  
weight  of  plant.  In  fact,  photosynthesis  rate  and  fresh  weight  
efficiency   and   electric   power   consumption.   Production  
under   five   light   qualities   set   by   LED   system   were   highly  
efficiency   means   dividing   average   fresh   weight   by   electric  
correlated  (R2  =  0.80ሻ.  Fluorescent  lamps  had  wider  radiation  
power   consumption.   R   and   B   treatment   using   LED   systems  
range  than  LED  systems  had.  It  could  occur  that  plants  under  
had   much   lower   power   consumption   and   higher   production  
FL  treatment  had  big  fresh  weight  for  photosynthesis  rate.    
efficiency  compared  to  FL.  Production  efficiency  of  B,  which  
Dry   weight   had   big   difference   among   individuals.   Figure   6   was   the   smallest   in   that   of   other   light   qualities   using   LED  
shows   the  relation  between  light  quality  and  dry   weight  rate   systems,   was  about  1.8  times  higher  than  that  of  FL.   One  of  
which   means   dividing   dry   weight   by   fresh   weight.   Dry   the   most   important   reasons   of   this   result   could   be   the  
weight   rate   had   small   difference   among   individuals.   Small   difference   of   energy   saving   ability   between   LEDs   and  
amount  of  dry  weight  rate  increased  in  FL  and  B  and  reduced   fluorescent   lamps.   On   the   other   hand,   the   large   difference  
in   R.   It   was   reported   that   lettuce   grown   under   red   light   had   among   light   qualities   using   LEDs   was   also   observed.  
the  smallest  dry  weight  among  that  under  monochromic  blue,   Production   efficiency   of   R   treatment   was   about   1.7   times  
blue-­green,   green   and   red   light   (Hirai   et   al.   2006).   In   our   higher  than  that  of  B  treatment.  This  difference  was  because  
research,  plant  in  R  treatment  had  as  big  dry  weight  as  that  in   red   light   has   lower   power   consumption   than   blue   light.  
B   treatment   had.   This   result   was   different   from   previous   However,   power   consumption   differs   from   light   sources,   so  
research,  but  dry  weight  rate  was  much  lower  in  R.     this  data  is  one  of  examples.  
5 4.  CONCLUSIONS  
Dry weight rate (%)

4 In   conclusion,   monochromic   red   light   might   be   the   most  

effective   for   photosynthesis   and   growth.   However,  
3 monochromic  light  also  makes  plant  shape  bad  and  the  plant  
is  worthless  as  a  product.  To  improve  plant  shape,  plant  must  
2 be  radiated  mixture  light.    
1 REFERENCES  
0 Akoyunoglou,   G.   and   Anni,   H.   (1984).   Blue   light   effect   on  
chloroplast   development   in   higher   plants.   In   Senger   H  
R RB SL-­L FL-­L B FL
  (ed.),   Blue   light   effects   in   biological   systems,   397-­406.  
Springer-­Verlag,  Berlin.  
Fig.   6.   Relation   between   light   quality   and   dry   weight   rate:   Balegh   SE.   and   Biddulph   O.   (1970).   The   Photosynthetic  
Light  quality  contains  more  red  light  in  order  from  left.  Each   Action  Spectrum  of  the  Bean  Plant.  Plant  Physiol.,  46,  1-­
value  is  the  mean  ±  standard  errors  of  replications.   5    
  Björkman  O.  (1973).  Comparative  studies  on  photosynthesis  
in  higher  plants.  Photophysiology,  8,  1-­63  

         
    
 
Flint  and  McAllister.  (1937).  Wavelengths  of  radiation  in  the   Mori  Y.  and  Takatsuji  M.  (2000).  Influence  of  laser  and  LED  
visible   spectrum   promoting   the   germination   of   light-­ Lights   on   the   growth   of   lettuce.   The   Illuminating  
sensitive   lettuce   seed.   Smithsonian   Inst.   Publs.,   MIisc.   Engineering  Institute  of  Japan,  33,  283-­284.  
Collections,  96,  1-­8.   Mori   Y.,   Takatsuji   M.   and   Yasuoka   T.   (2002).   Effects   of  
Folta.  (2004).  Green  Light  Stimulates  Early  Stem  Elongation,   Pulsed   White   LED   Light   on   the   Growth   of   Lettuce.  
Antagonizing   Light-­Mediated   Growth   Inhibition.   Plant   J.SHITA,  14  (3),  136-­140.  
Physiology,  135,  1407-­1416.   Takemiya   A.,   Takahashi   Y.   and   Shimazaki   K.   (2007).   Leaf  
Folta.   and   Maruhnich.   (2007)   Green   light:   a   signal   to   slow   Temperature   Reduction   by   Blue   Light-­dependent  
down   or   stop.   Journal   of   Experimental   Botany,   58   (12)   Stomatal  Opening.  Cryobiology  and  Cryotechnology,  53  
3099-­3111.   (1),  1-­5.  
Hirai   T.,   Amaki   W.,   Watanabe   H.   (2006).   Effects   of   Terashima   I.,   Fujita   T.,   Inoue   T.,   Wah   S.C.   and   Oguchi   R.  
Monochromatic   Light   Irradiation   by   LED   on   the   (2009).   Green   Light   Drives   Leaf   Photosynthesis   More  
Internodal   Stem   Elongation   of   Seedlings   in   Eggplant,   Efficiently   than   Red   Light   in   Strong   White   Light:  
Leaf  Lettuce  and  Sun  flower.  J.SHITA,  18  (2),  160-­166.   Revisiting   the   Enigmatic   Question   of   Why   Leaves   are  
Green.  Plant  Cell  Physiol.,  50  (4),  684-­697