Phyto Tron

THE BOTANICAL REVIEW
VOL. 46 OCTOBER-DECEMBER, 1980 No. 4
PHYTOTRONS
R. J. DOWNS
Director o f Phytotron
North Carolina State University
Raleigh, North Carolina 27650
Abstract ........................................................................................... 447

Introduction ....................................................................................... 450
Phytotron Design ................................................................................. 455
Greenhouses ................................................................................. 460
Controlled Environment Chambers ....................................................... 462
Air Flow ................................................................................. 462
Temperature ............................................................................. 463
Light ..................................................................................... 464
Humidity ................................................................................. 464
Control Systems ....................................................................... 465
The Phytotron Building ..................................................................... 466
Special Facilities ............................................................................. 466
Phytotron Size ............................................................................... 470
General Design Criteria ..................................................................... 471
Design Problems ............................................................................. 471
Phytotron Organization ........................................................................... 475
Criticisms of Phytotrons ......................................................................... 476
Phytotron Research ............................................................................... 481
Biological and Technical Service ......................................................... 481
Advantages of Phytotrons ....................................................................... 483
Environmental Measurements ............................................................. 483
Efficient Maintenance and Operation ..................................................... 484
Biological Aspects ........................................................................... 484
Acknowledgments ................................................................................. 485
Literature Cited ................................................................................... 485
ABSTRACT
Phytotrons are the most complex form of controlled-environment fa-

cility. Artificially and naturally lighted controlled-environment rooms and
cabinets are used in conjunction with incubators, seed germination cham-
bers, roomettes, photoperiod rooms and mechanically-refrigerated green-
houses to provide a multiplicity of environmental conditions. Phytotrons
are distinguished from the installation with a few plant growth chambers
The BotanicalReview 46: 447--489, October-December, 1980.

9 1980The New York Botanical Garden
447
448 THE BOTANICAL REVIEW
by the fact that phytotrons are operated in such a way that a wide range
of several environmental factors can be studied simultaneously. Phyto-
trons have an operating staff of specialists to maintain the system and the
experimental material. Scientists, therefore, concentrate on research
rather than maintenance and operation of equipment. Phytotrons also
make efficient use of controlled environment space, since they receive
steady use without periods of inactivity and the downtime due to mal-
functions that often characterize plant growth chambers.
Although few, if any, technical problems in phytotron construction
exist today, phytotron design suffers from the same lack of information
that restricts performance of plant growth chambers; such as inadequate
data on the physical characteristics of lighting systems and relative hu-
midity control; especially those physical parameters that are dependent
on, and influenced by, the biological material or by other environmental
factors.
Many excellent phytotrons have been constructed. Some, like those in
Australia, France, New Zealand, and North Carolina, are general purpose
phytotrons. Others are designed for particular plant species such as the
rice research facility in the Philippines, or as in the USSR and Hungary,
for specific research objectives like cold hardiness. The value of phyto-
trons in the general strategy of biological research is exemplified by the
fact that one or more phytotrons ranging from 100 to over 600 m2 can be
found in at least 19 countries. The latest of these facilities are located in
Canada and the USSR.
Phytotrons are used primarily to investigate how environment controls
and modifies plant growth and development, but they are used also to
complement and supplement field and greenhouse research in areas like
plant breeding and introduction of new plant species and varieties. Syn-
ecology is built on a foundation of autecology and the key word in the
definition of autecology is environment. Phytotrons, therefore, also play
an important role in many phases of ecological research. Since plants can
be grown and developed in phytotrons at rates, and with chemical com-
positions, that match definitions of "normal" they allow detailed study
of the physiological and biochemical systems affected by climatic stress.
Phytotrons are an efficient method of managing controlled-environment
facilities, whether they are used as plant growth chambers to provide
constant, reproducible conditions for biochemical studies or for research
like simulation modeling which virtually requires simultaneous use of a
wide range of several environmental factors. Despite the efficiency of
operation and regardless of the manner in which they are used, however,
the scientific results still depend on the quality of the investigators doing
the research.
PHYTOTRONS 449
RESUMEN
Los fitotrones son la forma mas completa de "ambiente controlado"

disponible. Cuartos y gabinetes de "ambiente controlado," artificial y
naturalmente iluminados, son usados conjuntamente con incubadoras,
c~imaras para la germinaci6n de semillas, cuartos para el estudio de la
duraci6n del fotoperiodo, e invernaderos mecfinicamente refrigerados
para proveer las facilidades mas completas de "ambiente controlado."
Los fititrones se distinguen de una simple instalaci6n de cfimaras para el
cultivo o crecimiento de plantas, por la condici6n de que son operados
de tal manera que permiten que un gran rango de factores ambientales
puedan ser estudiados simult~ineamente. Los fitotrones poseen personal
especialisado en el mantenimiento de todo el equipo, y material para
experimentos necesario para cualquier tipo de estudio. De 6sta manera,
los cientificos pueden concentrarse en el estudio y no en el cuidado y
mantenimiento del equipo usado. Los fitotrones hacen uso continuo y
eficiente de todas sus facilidades al no tener ningun periodo de inacti-
vidad o debido a real funcionamiento que periodicamente caracteriza las
cfimaras de crecimiento de plantas.
De todas manera, algunos, si los hay, problemas t6cnicos en la con-
struccion de fitotron y disefio, se deben a la falta de informaci6n que
restringe el uso de las c/~maras de cultivo de plantas, como es la infor-
maci6n inadecuada de las caracteristicas fisicas del sistema de luz y los
controles de humedad relativa, especialmente los par/tmetros fisicos que
dependen o son influenciados por condiciones biol6gicas u otros factores
ambientales.
Excelentes fitotrones han sido construidos, algunos como los de Aus-
tralia, Francia, Nueva Zelandia, y Carolina del Norte fueron hechos para
uso general. Otros fueron disefiados para especies particulares de plantas
como por ejemplo en las Filipinas, para el estudio del arroz, o como el
de USSR y Hungria para un objetivo especifico, como es el estudio de
la resistencia a baja temperaturas en plantas. E1 valor de fitotr6n en el
estudio biol6gico, lo demuestra el hecho de que uno o m/ts fitotrones con
un espacio de 100 hasta m~ts de 600 mz estfin en uso en por 1o menos 19
paises. Los tiltimos de 6stos fueron construidos en Canadfi y USSR.
Los fitotrones son usados principalmente para el estudio de como el
"ambiente controlado" modifica o cambia el desarrolo y crecimiento de
la planta, pero tambi6n son usados para complementar o suplementar
experimentaci6n en invernaderos o el campo, en estudios sobre repro-
ducci6n y en la introducci6n de nuevas variedades y especies. La sine-
cologia se basa en autecologia y la principal palabra en la definici6n de
autecologia es "ambiente." Los fitotrones juegan un rol importantisimo
en varias fases de la experimentaci6n ecol6gica. Ya que las plantas pue-

den cultivarse y crecer en proporciones exactas, con proporciones qui-
micas que son id6nticas a las "normales," los fitotrones permiten el es-
tudio detallado de los sistemas fisiol6gicos y bioquimicos los cuales son
afectados por las presiones ambientales.
Los fitotrones proveen un m6todo eficiente de manejar facilidades de
ambiente controlado no obstante si son usados como cfimaras de creci-
miento, para mantener condiciones ambientales fijas (constantes) para
estudio bioquimico o para experimentaci6n en la imitaci6n de la natu-
raleza, que virtualmente requiere usos simult~neos de una gran variedad
de condiciones ambientales. Atin teniendo en cuenta la eficiencia o la
operaci6n o la manera como se usa, de todos maneras los resultados
cientificos dependen de la habilidad del estudiante o la persona que lo
efectfia.
"Environment seems to be the key word in an amazing number

of unsolved or partially solved problems in biology." (Went)
INTRODUCTION
The term phytotron was first applied to the Earhart Laboratory at the
California Institute of Technology (98). The combining forms, p h y t o =
plants and -tron -- instrument, were used to compare the complexity of
the Earhart Laboratory to that of the betatrons and cyclotrons being
constructed by physicists at the time. The p h y t o t r o n designation has
since been applied to any comprehensive set of controlled-environment
chambers and glasshouses used to study the effects of light, temperature,
humidity, and other environmental factors on plant growth and devel-
opment. A proper definition of a phytotron, however, would include that
the facility is operated in such a way that a wide range of environmental
factors can be studied simultaneously. It follows that a phytotron is op-
erated by a management team, consisting of scientists experienced with
research on whole plants, assisted by mechanical and electronic techni-
cians with special knowledge of environment control problems. Thus
three or four plant growth chambers used one or two at a time, while
being maintained on the fix-it-when-it-breaks principle, does not consti-
tute a phytotron.
Phytotrons arose in the natural sequence of events resulting from the
need to control the environment during the course of plant research pro-
grams. Biologists have always been aware of the advantages of carrying
out experiments under rigidly controlled conditions, but with the equip-
ment available they were rarely able to do so. In fact, many experiments
have been conducted under conditions so poorly defined as to be non-
PHYTOTRONS 451
reproducible. Since no amount of statistical manipulation can substitute

for lack of knowledge of the experimental conditions, results have often
been contradictory. For example, Briggs et al. (12) reviewed experiments
on the effects of electric fields on plant growth that had been conducted
over a period of 19 years and concluded that uncontrolled, and conse-
quently undefined, environmental factors vitiated most of the results.
Weed control, irrigation and fertilizer applications in the field supply
a small degree of environment control that is adequate for many kinds of
research. Other investigations can proceed quite well in glasshouse struc-
tures. Many problems in plant science, however, like the research with
electric fields, cannot be solved under these conditions because naturally
fluctuating environmental events confound the experimental results. In
glasshouses, low light levels during winter often negate the advantage of
temperature control and the light available during summer has to be dras-
tically reduced by shading in order to keep the temperatures low enough
for plant survival. Thus it is not surprising that glasshouse experiments
conducted in the fall often produce results different from those carried
out in the spring and summer.
Realization by biologists of how little they understood the way in which
various environmental factors affected the many physiological processes
influencing plant growth and development motivated them to develop
devices in which plants could be grown under more controlled conditions.
Some plant scientists attempted to obtain precise temperature and hu-
midity control by building small, naturally-lighted chambers inside
glasshouses (47). Others constructed chambers that used high wattage,
nitrogen-filled, incandescent-filament lamps, often called Mazda lamps,
as a light source (19, 39, 85). Although a few of these early systems, such
as Tottingham's (89), controlled temperature by circulating mechanically-
refrigerated brine through pipes around the chamber walls, many others
used outside air for temperature control and could only be used effec-
tively during the winter months.
In the early 1920's air-conditioning was rarely encountered and even
by 1933 only a dozen or so office buildings, and perhaps 20 department
stores, in the United States were air-conditioned. The use of mechanical
refrigeration, however, was increasing rapidly. Air-conditioned theaters
increased from two or three in 1922 to better than 300 by 1933 and over
four million electric refrigerators could be found in homes, stores and
restaurants (43). The desirability of adapting mechanical refrigeration to
temperature and humidity control in plant growth rooms seems so ob-
vious that it is difficult to understand why so few plant scientists used
the systems. Equipment for single chamber use was available in sizes
ranging from fractional tonnage units characteristic of domestic refrig-
erators to larger compressors that operated with direct expansion air
handlers of up to three tons capacity. Still larger systems could have been
obtained for multiple chamber designs by adapting the methods used for
ice manufacture. A system of this latter type was used to control tem-
peratures in the early plant growth chambers constructed at the Boyce
Thompson Institute (3, 4). About 1937, four chambers, originally designed
as part of a winery at the Plant Industry Station at Beltsville, Md., were
modified for plant growth chamber use. Like the rooms at the Boyce
Thompson Institute, temperature was controlled by circulating calcium
chloride brine chilled by ammonia compressors to individually-con-
trolled, liquid-to-air heat exchangers. The Beltsville rooms were often
used to provide a range of environmental conditions, so the facility could
be considered a small phytotron. However, the first facility designed from
the outset to operate as a phytotron was located at the Kaiser Wilhelm
Institute in BerlinoDahlem in 1938 (101). Temperatures were controllable
over a range of 0 to 40~ in four artificially-lighted rooms and one dark-
room. Humidity reportedly could be controlled from 20 to 80%; although
from the description of the facility, humidities below 50% seem unlikely.
Considering that a few plant growth rooms with an excellent degree of
temperature control were built during the 1930's, some factor other than
availability of suitable mechanical refrigeration systems must have been
inhibiting more widespread development of controlled-environment
rooms. Since completely controlled plant growth rooms must rely on an
artificial source of illumination to produce the relatively large amount of
light necessary for obtaining uniform conditions throughout the year, the
major obstacle to the early development of the plant growth chambers
was undoubtedly the lack of satisfactory light sources. Many of the early
chambers, like those at the Boyce Thompson and the Kaiser Wilhelm
Institute, used incandescent lamps separated from the growing area by
a transparent barrier containing a thermal filter, typically water, to re-
move the large amounts of radiant heat emitted by the lamps. The irra-
diance at plant level was not very high, because the incandescent lamp
is an inefficient light source. Moreover, many plants reacted adversely
to the spectral distribution of illumination from incandescent lamps.
The chambers at Beltsville used 75 ampere, alternating current, carbon
arc lamps for plant growth lighting (Fig. 1). The irradiance was sufficient-
ly high for good plant growth but the spectral energy distribution was
unsatisfactory until a small amount of light from incandescent lamps was
added. The improvement in plant growth was much greater than could
be accounted for by the small increase in illumination and it was con-
cluded that the incandescent lamps were balancing the predominantly
blue emission of the carbon arc by building up the red end of the spectrum
(71). Although the carbon arc plus incandescent lamps proved to be an
excellent light source for plant growth and was used almost continuously
PHYTOTRONS 453
Fig. I. Carbon arc lighted controlled-environment room at Beltsville that provided over
20,000 lux at pot level.
for over 30 years, the system required constant maintenance and great
care had to be exerted to remove phytotoxic gases and to prevent the
short wavelength ultraviolet radiation generated by the arc from reaching
the plants.
Plant growth chambers did not become a practical research tool until
fluorescent lamps became generally available commercially. As soon as
these lamps could be obtained, reports began to appear showing that the
spectral emission of the commercially-produced lamps was very suitable
for plant growth (35, 38, 64). The high output of visible light per watt of
power, combined with low levels of radiant heat, made the lamp espe-
cially useful in controlled-environment room applications. Moreover, the
broad source aspect of the fluorescent tube, as contrasted with the point
source configuration of incandescent lamps, made it easier to achieve a
high degree of uniformity of illumination over the plant growing surface.
Flux densities from these early, low current, lamps were inadequate but
the rapid development of the higher output, eight-foot slimline lamp al-
lowed biologists to obtain light levels sufficient for satisfactory growth of
a large number of species (7, 13, 72) (Fig. 2). By 1960 the advent of highly
loaded 1500 milliamperes (ma) Power Groove and VHO fluorescent lamps
enabled researchers to build chambers in which an illuminance of 40,000
Fig. 2. Early, 1947, controlled-environment room equipped with slimline fluorescent

lamps to provide an illuminance of 15,000 lux at bench level.
to 50,000 lux could be maintained at plant level (Fig. 3). Thus the avail-
ability of reliable air conditioning systems, low radiant heat, biologically-
efficient, fluorescent lamps, and the interest and advice of lamp company
engineers (notably General Electric's Joe Ditchman) stimulated the con-
struction of plant growth chambers. Research objectives were obviously
somewhat restricted when only one or two plant growth chambers were
available. The phytotron idea was developed as a means of easing these
restrictions.
The phytotron concept proposed by F. W. Went became a reality large-
ly because of the support of H. B. Earhart and the foresight of R. A.
Millikan, President of the California Institute of Technology. Upon its
completion in 1949, numerous scientists, many from other countries, vis-
ited the facility to observe and use the wide range of precisely controlled
environmental conditions. Thus the phytotron concept was disseminated
worldwide, and soon phytotrons of various sizes were under construction
in many other countries (Table I). Although the Earhart Laboratory in-
spired construction of phytotrons throughout the world, additional ones
were not built in the U.S.A. for nearly 20 years. About 1967 the Biotron
in Wisconsin was put into operation and shortly thereafter the South-
eastern Plant Environment Laboratories (SEPEL) with phytotrons in
PHYTOTRONS 455
Fig. 3. Controlled-environment room constructed in 1963 at Beltsville using highly load-

ed Power Groove lamps to obtain an illuminance of 40,000 lux at bench level.
Durham and Raleigh were opened in North Carolina. Virtually all phy-
totrons result from the dedication of one or two individuals who are rarely
identifiable after the facility becomes operational. The SEPEL phyto-
trons, for example, resulted from the combined efforts of P. J. Kramer
of Duke and K. R. Keller at North Carolina State University, and the
facility at Gif-sur-Yvette, France was built because of the endeavors of
Paul Chouard.
PHYTOTRON DESIGN
A phytotron is often visualized as consisting of a combination of tem-

perature-controlled glasshouses and walk-in, artificially-lighted plant
growth chambers because the original phytotron was constructed in this
manner. Many phytotrons, however, do not follow this plan. Those in
Belgium (10), Hungary (74, 75) and Wisconsin (82) do not include any
glasshouses. In Australia at the CERES phytotron (62) artificially and
naturally-lighted reach-in cabinets are used in conjunction with temper-
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PHYTOTRONS 457
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Fig. 4. Cross section of Phytotron I. Biotron Institute in Japan. 1, glass room. 2, roof
spray. 3, air outlet. 4, air inlet. 5, air supply duct. 6, return air duct. 7, mixing chamber.
8, fresh air supply duct. 9, machinery room. 10, corridor. 11, detecting point. (Dimensions
are in meters.) (54)
ature and humidity controlled glasshouses and several phytotrons are

combinations of glasshouses and three or more sizes of controlled-envi-
ronment chambers (Table I). At Palmerston North (93) greenhouses and
slat houses are used as auxiliary equipment and the Oslo phytotron op-
erates in conjunction with a 3000 mz cultivation area and a 5000 m 2 broad-
leaf forest (65). Phytotron glasshouses are usually small, ranging from 9
mz (73) to 49 m 2 (99) although the phytotrons in France and the Biotron
Institute in Japan also operate super glasshouses of 1076 m z and 576 m2
respectively (17).
Phytotrons are usually designed in their entirety by architects and en-
gineers who are more familiar with comfort air conditioning than with
phytotrons. The input from biologists varies from a simple declaration of
the requirements of the research to be conducted in the facility to con-
tinuous involvement through all design phases. In only a very few in-
stances have key members of the phytotron staff shared design respon-
sibilities and followed the construction of the facility on a daily basis.
These examples, however, show that the greater the involvement of the
biologists and the phytotron operating personnel during planning and con-
struction, the more suitable the final results for the research programs
and the fewer the difficulties encountered in the daily operation.
The Designing Institute in Moscow apparently has designed, or at least
PHYTOTRONS 459
~" IA 2
i
Fig. 5. Cross section of phytotron II. Biotron Institute in Japan. 1, glass room. 2, roof
spray. 3, corridor. 4, machinery room. 5, mixing chamber with air washer. 6, air supply
duct. 7, air outlet. 8, air inlet. 9, return air duct. 10, fresh air supply duct. 11, detecting
point. 12, movable ladder. (Dimensions are in meters.) (54)
advised on the development of new phytotrons at Odessa, Mironovka,

and Krasnodar; including the controlled-environment facilities (2). The
trend, however, during the past ten years has been to simply construct
a building to house commercially-produced controlled-environment
chambers. Koito chambers, for example, are used at IRRI (106), and
Conviron units make up the phytotrons at Martonvasar (75) and Leth-
bridge (37), and EGC chambers are used in SEPEL; although the latter
have been considerably modified at the Raleigh phytotron.
The advantage of using commercially-manufactured chambers is that
the performance characteristics have been thoroughly documented. Re-
search, therefore, can begin immediately without any substantial period
of time lost in testing and redesigning. This advantage is forfeited, of
course, when the required environmental control or chamber dimensions
are outside the range normally produced by the manufacturer. Since per-
formance is known in advance, the objection to the use of prefabricated
chambers is largely centered around the idea that they are not reliable
and that maintenance is difficult. This concept developed from personal
communication among biologists telling of the many operational problems
encountered with commercially-manufactured chambers. Reports of sat-
isfactory operation, on the other hand, are rarely disseminated. Certainly
maintenance of many prefabricated chambers is much more difficult than
is really necessary. The poor reliability reputation, however, is question-
able. For example, a stock commercial chamber recently tested at the
Southeastern Plant Environment Laboratories (SEPEL) has run contin-
C11 C12 C13
C21 C22 C23

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C31 , 32 I C33 (cm)
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Rhizosphere
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Fig. 6a. Measuring points of humidity in plant population. (58)
uously for over two years with only one minor malfunction. The impli-
cation is that if prefabricated chambers are installed correctly at the out-
set and maintained properly, reliability is reasonably good.
The decision of whether or not to use prefabricated chambers thus
seems to depend on two factors. First, does the proven environmental
control capability of commercially-manufactured chambers fulfill the re-
quirements imposed by the research strategy. Second, does the designing
engineer have sufficient experience with environment control for biology
to develop better systems than those available commercially. Unfortu-
nately the latter is rarely true.
Greenhouses
Phytotron glasshouse designs are very similar. Major differences are
the use of water flowing over the roof and the direction of air flow. The
first glasshouses built at the Biotron Institute in Japan introduced the
PHYTOTRONS 461
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r
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I~ I~ \ \ ".. \ ~ -- - __ ----- j I I I
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a21"r "ra22 ~a23
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_t_
Outlet
Fig. 6h. Vertical section of humidity distribution in c u c u m b e r plant population in up-
ward air flow of 0.2 m sec -I u n d e r air temperature of 25~ and relative humidity of 50% in
a phytotron glass room. (Arrows indicate the direction o f air flow.) (58)
conditioned air near the top of the structure and circulated it downwards
over the plants (Fig. 4). In later units (Fig. 5) the air moved upward
through the floor in the manner of the Earhart Laboratory. The more
recent structures also have a system of running water over the roof. The
film of water on the roof is too thin to function effectively as a thermal
filter, but since it is evaporatively cooled by spraying through nozzles
the water does remove the heat absorbed by the glazing and structural
members.
Unique designs can be found at Stockholm and at CERES. At the
Stockholm phytotron, glasshouse heat loss on cloudy days and at night
can be drastically reduced by automatically-operated darkening devices
on the glass walls. A lamp bank can be moved over the inner glasshouse
roof to provide plant growth room level artificial lighting (100). In Aus-
tralia, a reverse cycle refrigeration system is used for air-conditioning the
glasshouses. Excess heat is transferred to a thermal storage pond during
the day and recycled to the glasshouses at night. Heat from the lamp lofts
of the artificially-lighted chambers is used to help balance the heating
Outlet Inlet
I I
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r ~ I i ~ l "
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l I I I J \
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Fig. 6e. Vertical section of humidity distribution in cucumber plant population in lateral
air flow of 0.2 m sec-1 under air temperature of 25~ relative humidity 50% and artificial
light in a growth cabinet. (Arrows indicate the direction of air flow.)
demand. Hot water is produced by solar heating panels located on the

roof (68).
Controlled Environment Chambers

Airflow.---Air may move across the room (1, 23, 81) upwards through
the floor in France (66) and Oslo (65), or in reverse from top to bottom (30,
62, 93). The supposed merits of the different air flow directions are sup-
ported by few comparative data. Air flowing across the room would seem
to result in a temperature gradient over the plant growing area, especially
if light levels are high, but Forrester (34) claims that plants placed in
these rooms create an air turbulence that reduces the gradient to a neg-
ligible level. The rationale for upwards movement through the floor is
that a laminar flow can be obtained. Unfortunately the laminar flow is no
longer present when plants are placed in the room. This results in a high
air velocity between the plants and a low velocity above the plants where
the radiation effects are greatest (45). Another difficulty with upward air
flow is that a solid floor must be constructed under the perforated or
slotted one. Water, plant, and substrate debris fall into the plenum and
if it is not removed frequently it interferes with humidity control and
becomes a source of saprophytic contaminants and pathogenic infections.
Recently Matsui et al. (58) measured humidity distribution produced with
PHYTOTRONS 463
I" / / / .i - - ~ ~X \ \
J / ~\ \ \ \
a )a22 23
I Rhizosphere I
Inlet
Fig. 6d. Vertical section of humidity distribution in cucumber plant population in down-
ward air flow of 0.2 m sec -~ under air temperature of 25~ relative humidity of 50% and
artificial light in a growth cabinet. (Arrows indicate the direction of air flow.)
each air flow system (Figs. 6a, b, c, d) and concluded downward, often
called reverse, air flow was the most reliable.
Temperature.--The temperature range usually available in phytotrons
is from 5 to 35 or 40~ although a number of facilities have a few rooms
that can provide - 1 0 or, as at Ostankino, as low as -60~ (21). Tem-
peratures below about 5~ cannot be maintained without some method
of defrosting the cooling coils. The deicing process usually leads to a rise
in temperature of a magnitude and duration commensurate with the de-
sign. Although the importance of deicing design is obvious, very little
information on the subject has been published.
Higher than average maximum temperatures, up to 50~ are provided
at CERES (62), Palmerston North (93), Irhoutsk (2), and Shanghai (105).
In practice phytotrons that indicate maximum temperatures of 35~ can
provide higher ones but do not do so because the high temperatures
shorten the life of electrical components. Fan life, for example, will be
reduced 42% at 50~ as compared to those operating at 25~ Thus con-
trolled-environment rooms that operate above 35~ for an appreciable
time have special electrical systems designed for high temperature op-
eration.
464 T H E B O T A N I C A L REVIEW
Most phytotrons control air temperature. Radiation effects are elimi-

nated by placing the temperature sensors in aspirated housings where
they are shielded from the light source. Although plant temperature is
rarely known in the field or in glasshouses, some researchers insist that
it be known and regulated in controlled-environment chambers. Usually
this is not possible except in darkness, because the radiation from the
light source does not strike all portions of the plant equally. Moreover
most temperature sensors also measure radiant energy to a degree that
depends on the material, size and color of the sensors and the way they
are made. Remote sensing devices that measure temperature by detecting
infrared emitted by the leaf can also measure reflected long wavelength
radiation that adds an artificial increment to the true plant temperature.
The temperature of a single leaf, however, can be measured and Matsui
and Eguchi (56) have developed methods of using leaf temperature as a
feedback signal to the air temperature controls to maintain the desired
value.
Light.--The majority of the artificial light systems used in phytotrons
are composed of fluorescent and incandescent lamps. Only fluorescent
lamps are used at Oslo (65). Occasionally a few high intensity discharge
lamps (HID) are added (104) to the fluorescent system and some phyto-
trons use HID lamps alone or in combination with incandescent (1, 11,
55, 83, 94) or fluorescent lamps (81). Light levels range from 1000 to
100,000 lux; the latter obtainable only in a few rooms equipped with
Xenon arc lamps. Plant growth chambers in the phytotron at Shanghai
are lighted with Xenon arc lamps but the illuminance is only about 30,000
lux because the lamps are a considerable distance from the plants (P. J.
Kramer, personal communication).
Light sources are usually separated from the plant growing area by a
transparent barrier. Most installations using HID lamps (60, 95) include
water baths in the barrier to serve as thermal filters (Fig. 7). These water
baths, however, have to be carefully designed in the manner of the sys-
tems at Palmerston North (48). The importance of this seems to be lost
on chamber designers in the U.S., where HID lighted rooms are being
constructed with only a single glazed barrier of unknown thermal char-
acteristics.
Humidity.--Precise humidity control is costly and difficult, especially
below 50%. To maintain low humidity, water and nutrient solution must
be added automatically without spillage and the chamber door opened
only if some malfunction occurs. Solution draining from the plant con-
tainers must be collected and removed without spillage. Below 50% RH,
chemical driers have to be used in addition to the cold plate dehumidifier.
An important fact about chemical driers that often seems to be over-
PHYTOTRONS 465
Fig. 7. Metal halide and incandescent lighting system in operating position above a
glass-water thermal barrier at the Climate Laboratory, Palmerston North, New Zealand. (93)
looked is that their primary application is to provide low relative humid-

ities at low temperatures. Thus the efficiency of chemical drying de-
creases for air entering conditions above 10~ (97). Controlling relative
humidity above 50% is relatively a much easier task. Therefore, humidity
control in most of the controlled-environment rooms of phytotrons covers
the range of 50 to 90%, with a few chambers that can provide 100% and
a few in which humidity may be maintained as low as 15%.
Control systerns.--Most phytotrons have access to computer systems
for analysis of experimental results. Several phytotrons, such as SEPEL
and Palmerston North, employ microprocessor-based, programmable
controllers, but except for a few facilities like the Wisconsin Biotron and
the phytotron at Oslo, phytotrons do not usually use a computer for
control or for acquisition of data concerning environmental parameters.
The most elaborate use of computer control is at the Biotron Institute in
Japan where biological inputs are used to regulate the environmental
variables. Nondestructive digital image processing, for example, is used
to obtain plant growth information for feedback to adjust the environ-
mental conditions during the computer control of the growth processes
(57).
The Phytotron Building

The structure housing the controlled-environment chambers must be
designed around the traffic patterns generated by the research objectives
of the facility. Floor plans of the various phytotrons, however, show
many similarities (Figs. 8a, b). Most phytotrons are planned to allow
unrestricted movement of research material between controlled-environ-
ment chambers and to special facilities and treatment rooms, even where
there is no intention of moving plants on a daily basis. Conversely, de-
spite the obvious advantages, few phytotrons have methods o f entering
the controlled environment spaces during the dark period without allow-
ing light to enter also.
Phytotron buildings are usually designed as limited access structures.
Clean-room type plans, however, have not proven practical and few phy-
totrons set up originally with stringent clean-up procedures have kept
them in use. Nevertheless, insect infestations in phytotrons can virtually
always be traced to personnel bringing insects into the facility on their
persons or on plant material. Some kind of clean-up procedure is ob-
viously necessary, and at CERES and SEPEL a practical compromise
was adopted that requires researchers and visitors to wear coveralls and
shoe covers. Personnel also are asked to wash their hands and comb their
hair before donning the protective covering. All supplies, equipment, and
especially plant material should be fumigated before bringing it into the
clean area and several phytotrons contain gas-tight rooms especially for
this purpose. Additional contaminant protection is afforded by the fact
that phytotron buildings are usually somewhat pressurized. Air entering
the building is filtered and the air returning for mixing and recirculation
is refiltered. Filtration methods are adequate for removal of fungal spores
and some bacteria, and in many areas activated charcoal or final filters
are used to remove phytotoxic gases.
Special Facilities
Virtually every phytotron contains special equipment that reflect the
research objectives of the facility. Several phytotrons, for example, in-
clude dark rooms housing spectrographs (54, 60, 68, 81). These biological
spectrographs are usually grating instruments powered by carbon arc or
Xenon arc lamps. The spectrum produced may be 270 • 18 cm or more
and cover a range of 350 to 800 nm. Energy levels may be as high as 3.45
W m-2 nm -~. In addition to the biological spectrograph, the phytotron at
Tsukuba, Japan contains a Xenon arc lamp powered monochromatic light
room (81) and the Biotron Institute has developed chambers in which the
total and spectral irradiance can be controlled and programmed (59).
A wind tunnel was part of the Earhart Laboratory (98) and similar
PHYTOTRONS 467
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PHYTOTRONS 469
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Fig. 9. Schematic of the CSTR system gas-sampling unit for the Phytotron. Note: All
sampling lines and the sampling manifold are insulated and heat-taped.
systems have been included in several later phytotrons (54, 55, 60, 82).
A lighted wind tunnel using temperature and humidity-controlled air is
planned for the new phytotron at Odessa (2).
Continuously stirred tank reactors (Fig. 9) for exposing plants to gaseous
air contaminants were installed at SEPEL (40). These systems use a
dynamic, negative pressure, single-pass air flow for maintaining plants in
a uniformly mixed exposure atmosphere. Inlet air is temperature- and
humidity-controlled and each unit is equipped with an HID metal halide
lamp that provides 35 klx at plant level. Rates of gas uptake, net photo-
synthesis and transpiration, can be obtained by monitoring inlet and out-
let gas streams. A refined version of the CSTR system was developed for
precision measurements of gas uptake by whole plants (78).
Root temperature control is provided in special facilities at a number
of phytotrons, such as CERES (68), The Biotron Institute (55), and SE-
PEL. In the Biotron Institute system, containers of soil or other substrate
are immersed in a water bath that allows soil temperatures 5~ higher or
lower than the air temperature. A larger root to air temperature differ-
ential cannot be used satisfactorily because uniformity of temperature
throughout the root zone in sand and soil cultures is difficult to achieve
if root and aerial temperatures are greatly divergent (92). The temperature
gradients in the water bath method, however, can be used in some studies
to simulate naturally-occurring soil gradients (79). Uniform root environ-
ments are obtained at SEPEL by growing plants in a continuous flow
liquid culture with automatic pH and temperature control (69). The con-
tinuously flowing nutrient solution avoids the temperature gradient prob-
lems encountered with sand and soil substrates, so root zone tempera-
tures can be maintained between 10 and 35~ at any ambient temperature
over the same range.
Frost rooms are unique facilities located at CERES (5) and Palmerston
North (77). White frosts are formed by keeping the relative humidity at
100% while the temperature is below 0~ while black frosts are produced
with cold air flow of low moisture content. At Palmerston North, the
design has been refined so that rates of freezing and thawing, as well as
the duration and level of the low temperature period, can be controlled.
Since the root zone often does not freeze under naturally-occurring frost
conditions, plant containers are placed in insulated tanks in which the
temperature is controlled above chamber conditions by low voltage heat-
ing tapes.
Phytotron Size
Lang (53) suggested that a phytotron has a definite minimal and max-
imal critical mass. Certainly a phytotron must be large enough to supply
a significant number of environmental conditions. What constitutes a sig-
nificant number, however, depends on the research strategy of the facil-
ity. Thus three temperature and humidity controlled glasshouses and one
artificially-lighted plant growth chamber seem to serve as a phytotron at
Kagawa University (73), whereas at the Biotron Institute at Kyushu a
large number and great variety of chambers are required (Table II). Lang
(53) also suggested that when the number of individual chambers reaches
50 or more, the combinations of environmental factors become so large
that it is doubtful they can be used effectively. Many phytotrons, how-
ever, accommodate a number of investigators, not all of whom will be
interested in the same environmental variables or in the values selected
for a range of conditions. Thus to be research efficient, more than one
series of environmental conditions must be available. The SEPEL phy-
totron in Raleigh, for example, contains more than 50 chambers but the
demand for using the facility is so great that 10 more could be used with
ease. Nevertheless, it would seem that as phytotron size increases tech-
nological and research operations must eventually become unmanage-
able. With today's computer-oriented technology and the accumulated
experience at phytotron management, however, the number of units that
PHYTOTRONS 471
can be operated effectively is very large and the upper limit in size is
unlikely to be a factor to influence the design.
In practice the optimum size of a phytotron depends only on the type
of research to be conducted and the number of investigators that intend
to use the facility. Phytotron design, therefore, begins by determining the
research strategy of the facility and estimating the number of research
projects that will be accommodated simultaneously. Decisions can then
be made on the environmental factors to be controlled, those that will be
held constant, and the range of requirements of the various environmental
conditions.
Phytotron research strategies upon which the design is based may have
a single objective such as cold injury (104), or the studies may be of wide
scope but confined to a single species, genus or plant type such as rice
(106), tobacco (67) or trees (81). The general purpose phytotron is usually
the most complex because it is used for research on diverse problems
and many species are involved.
General Design Criteria

The most obvious design mandate for a phytotron is uniformity of
environmental conditions in the plant growing area. This requires the
smallest possible variation about the set point of each environmental
factor. For example, it would be realistic to expect temperature, mea-
sured with a thermocouple made with 0.51 mm wires mounted in a shield-
ed, aspirated housing, to vary no more than _+0.25~ if the chamber air
was conditioned by a secondary coolant system. As a general rule the
variation across the plant growing area in a well-designed chamber would
then be about double the variation at the set point sensor.
Less obvious design criteria are mechanical, electrical and electronic
components that exhibit a high degree of reliability and stability in a
design that provides maximum flexibility for future modification and mod-
ernization of equipment.
Design Problems
One of the major difficulties in designing a phytotron is that the biol-
ogist must, with rare exceptions, work with engineers and architects who
would be more comfortable designing an office building or elementary
school. As F. W. Went (16) pointed out, few air conditioning engineers
have any experience with controlling glasshouse and growth chamber
environments and the result is that a lot of unnecessary errors occur.
This is due in part to the fact that engineers seem to consider the design
of a phytotron a relatively simple, straightforward problem; although it
should be obvious that designing a phytotron is considerably more ar-
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PHYTOTRONS 473
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duous than providing lighting systems and air conditioning for an office
building. The phytotronist may have great difficulty in getting across the
idea that reliability of equipment must exceed what is usually acceptable
in conventional building design. The response to the high reliability re-
quirement is often the inclusion of back-up systems. Back-up systems for
everything would be a classic case of redundancy and would not be eco-
nomically feasible. Moreover, successful use of selectively placed back-
up systems requires an uncanny knowledge of which systems will have
the highest failure rate. Since in practice the required degree of reliability
is unlikely to be attained, much design effort should go into ease and
speed of maintenance. Unfortunately few engineers have any experience
with maintenance, so the probability of attaining an easily maintained
design is also low, unless assistance from experienced phytotronists and
physical plant engineers can be obtained during the early design stages.
The phytotronist may encounter equally insurmountable hardships in
trying to obtain enough flexibility in the design to permit modifications.
Several years may pass between the initial design and the actual use of
the facility. During that period, some research objectives may change
and new ones will surely be added. Consequently it is not unusual to
begin making modifications almost as soon as a phytotron is placed in
operation. Those accustomed to designing office buildings consider this
procedure a result of poor planning and thus avoidable, but this is not
necessarily true in research laboratories used as phytotrons.
Phytotron design is complicated by the fact that little if any research
is being conducted to determine optimum design parameters for con-
trolled environments for biology. As a result basic design is little different
than that used 15 years ago. Air flow and temperature interactions on
light output and lumen maintenance of highly-loaded fluorescent lamps,
packed closely together as they are in plant growth chambers, are not
understood. Air flow in aspirated housings for temperature and humidity
sensors is often too low. And we are still debating the proper air velocity
for controlled-environment rooms.
Although light sources have been separated from the growing area of
plant growth rooms for many years, the thermal characteristics of present
and potential barrier materials are poorly defined. The current trend to-
ward using high intensity discharge lamps as the main light source for
controlled-environment rooms calls attention to the fact that very little
has been published concerning the design of these lighting systems. Prac-
tically all the information concerning installation of HID lamps has come
from the Palmerston North phytotron (95).
Fluorescent-lighted controlled-environment rooms usually include in-
candescent lamps. Incandescent lamps are short-lived at best and when
used in controlled-environment chambers have a tendency to fail pre-
PHYTOTRONS 475
maturely (25). Moreover, incandescent lamps are inefficient light pro-

ducers and work should be in progress to find a biologically acceptable
substitute.
The amount of incandescent light most often used is 10% of the illu-
minance or 30% of the installed watts of the fluorescent system. These
values are estimates based upon the amount of incandescent light that
seemed satisfactory when added to a main light source of carbon arc (70)
or relatively low output fluorescent lamps (102). Although the optimum
fluorescent-incandescent ratio has never been determined, few existing
controlled-environment room designs allow for even a nominal increase
in wattage or number of incandescent lamps. Yet an increase may well
be desirable. Warrington et al. (95) note that incandescent lamps should
be added to metal halide HID lamp systems to procure the best plant
growth. Of particular importance is their recommendation that an HID-
incandescent ratio of 1:1 by installed watts should be used.
PHYTOTRON ORGANIZATION
Phytotrons may have no permanent research personnel other than the

directors, or they may operate around defined research groups attached
to the facility. At SEPEL, for example, phytotron users are either visiting
scientists or members of the various departments in the School of Agri-
culture and Life Sciences. Visiting scientists often use the phytotron di-
rectly but they may also operate through one of the SALS departments.
Research, therefore, reflects the programs of interest to the various de-
partments, although no restrictions are imposed and any scientist can
choose to conduct experiments on problems unrelated to ongoing re-
search. Many phytotrons operate in a similar manner. The IVT phytotron
at Wageningen is primarily a research tool for personnel in the Institute
of Horticultural Plant Breeding. The phytotron at the National Taiwan
University is mainly used by researchers in agronomy because of the
emphasis on rice and soybean investigations, but it is also available to
members of the other departments of the College of Agriculture.
The phytotron at Gif-sur-Yvette, in contrast, has a permanent staff of
about 40, divided into several research groups. Visiting scientists usually
work as part of one of these teams. The general objective of the Gif
phytotron is to study the physiology of morphogenesis of vascular plants,
in particular the physiology of reproduction (18). One research group
studies growth and development from the point of view of ecology.
Another is interested in the mechanisms that control and direct growth
of different kinds of meristems. Still other teams are investigating the
function of the various metabolic chains, such as the biosynthesis of
alkaloids. Thus fields of interest range from ecophysiological systems and
processes of morphogenesis to biochemical mechanisms and related bio-

physical reactions. These are the same fields of interest encountered at
other general purpose phytotrons such as SEPEL. The methods of ap-
proaching the research objectives, however, are different, because while
mission-oriented, team research is an efficient system it may not be prac-
ticable in some organizational structures.
The demand for space in phytotrons usually is so great that many of
them have review committees to evaluate the proposals for the use of the
controlled environment space. These committees meet at appropriate in-
tervals, monthly (27) to every four months (18), to evaluate the space
requests. Selection criteria are about as nebulous as those used by grant-
ing agencies for funding research. Will the research make a significant
contribution to the solution of a plant science problem? Is the scientific
quality of the proposed study obviously superior and competently orga-
nized? The phytotron committees, however, do not assume the role of
a power elite that decides on the kind of research to be conducted. The
committee members are experienced in the use of phytotron facilities and
act as advisors to the prospective users in developing a research design
that is compatible with and makes maximum use of the phototron equip-
ment. Visiting scientists usually receive special consideration and addi-
tional priority is given to problems of global concern over those of local
interest.
All true phytotrons are designed to investigate the effects and inter-
actions of a wide range of climatic conditions, usually on a specific bio-
logical response such as flowering, seed production, nutrition, protein
synthesis or disease resistance. Phytotrons are also called upon to pro-
vide a single set of conditions that serve as a reproducible standard en-
vironment. These standard conditions are, or should be, obtained from
experiments that establish the optimal or critical levels of the environ-
mental factors being controlled by using a wide range of several climatic
conditions simultaneously. Standard-environment type research gener-
ally receives a low priority for use of phytotron space and when the
environmental conditions are selected by intuition priorities are even
lower.
C R I T I C I S M S OF P H Y T O T R O N S
A major criticism of early phytotrons was the lack of flexibility in the

design, which made it extremely difficult to keep equipment updated and
to adapt to changes in research objectives. Some researchers object to
the Earhart method of moving plants from one room to another because
of assumed thigmomorphogenic effects, although experiments at Earhart
and later at CERES did not reveal any differences between plants that
PHYTOTRONS 477
were moved and those that were not (53). More recent studies (44, 88)
suggest that mechanical perturbations do have a detrimental effect on
plant growth. The possibility of detrimental effects from moving the
plants is often used to support the use of numerous reach-in cabinets
rather than fewer walk-in rooms. Phytotrons using only reach-in cabinets,
however, are limited by the fact that many species cannot be grown to
maturity. As a result many phytotrons employ both reach-in cabinets and
walk-in rooms. At SEPEL some of these chambers are under the control
of individual investigators and others are operated on the Earhart plan
where two or more researchers share the same space. Modern phyto-
trons, unlike early ones, are very flexible. Equipment can be modified
with relative ease and specialized facilities can be added to fit changes
in research objectives. The SEPEL phytotron, for example, has added
seed germinators, photoperiod rooms, air pollution exposure chambers,
special units for gas exchange studies and systems for root temperature
control.
Van Bavel and McCree (90) listed three points of criticism about con-
trolled-environment facilities in general. 1. Carbon dioxide levels are not
controlled. Operators of individually-owned plant growth chambers do
seem to rely on make-up air to maintain CO2 concentrations. The fallacy
of this approach can be easily demonstrated by simply measuring the
time course of CO2 concentration over a 24 hr period in a controlled-
environment chamber with an average load of plant material (Fig. 10).
Morse (61) calculated that 75% of the chamber air must be changed each
minute if make-up air systems are to prevent serious CO2 depletion. Since
very few make-up air systems are large enough to change this volume of
air, many growth chambers must be operated under CO2 stress conditions
(Fig. 11). Virtually all phytotrons, however, use some method of CO2
injection to provide CO2 control over a range from ambient to 1500 ppm
or more. Temporary, high levels of CO2 caused by workers entering the
controlled-environment chambers may not be controlled throughout the
phytotron. When necessary, worker breathing effects can be eliminated
by placing the affected chambers completely under automatic control and
by using respirators when the investigator must enter the room.
2. Light levels are too low, seldom exceeding 88 of direct sunlight.
Older fluorescent lighted rooms often produced a photon flux density of
200 IzE m -2 s-1 of photosynthetically active radiation (PAR) or less. This
was about equal to an illuminance of 15,000 lux and was inadequate for
proper growth and development of many plant species. Modern systems
using fluorescent and incandescent lamps, however, can produce a fairly
constant 700/zE m -z s-1 of PAR (27) and rooms using high intensity
discharge lamps produce 800-900/~E m -2 s-1 (94) which is enough to
provide a reasonable replica of a field phenotype (76).
478 THE BOTANICALREVIEW
550
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chamber filled with 4-week-old bean plants growing at a photon flux density of PAR of 670
/xE m-2 s-1.
Peak solar flux density of PAR is about 2000/~E m -2 s -1, although this
value varies somewhat with latitude and climate. A few phytotrons al-
ready have chambers where photon flux densities of solar magnitude can
be attained (82). Since the technical problems of obtaining very high light
levels are understood (95), any phytotron is capable of developing solar
flux density systems if the research strategy demands it. The merits of
solar light levels, however, remain to be proven experimentally. Jividen
et at. (46) failed to obtain plant growth commensurate with the investment
at light levels in excess of 750/zE m -z s -1. A more detailed study by
Warrington et al. (96) states that plants grown under high irradiance con-
ditions, 1665 /~E m -z s -1, were generally abnormal and not typical of
greenhouse or field grown material. Their results show that irradiance
levels equal to half full daylight are adequate for most studies.
3. A i r f l o w is too low. Air flow in the Earhart Laboratory was 0.16 m
s -1 and was undoubtedly too low. Most existing phytotrons, however,
provide air at 0.5 m s -1. This is based on the report of Morse and Evans
(62) which showed that air velocities in excess of this value reduced leaf
area and dry weight accumulation. Nevertheless Doorenbos (24) recom-
mends air flows of 1 m s -~ and van Bavel and McCree (90) suggest 1 to
3 m -1. These higher values seem to result from studies by Wadsworth
(91) that show no deleterious effect of air velocities as high as 1.5 m s -1.
PHYTOTRONS 479
Fig. 11. G r o w t h of B u s h Blue L a k e b e a n placed in a 2.44 x 3.66 m controlled-environ-

m e n t r o o m half filled with 1 m corn plants in which CO2 w a s not controlled (left), or con-
trolled at 350 p p m (right).
The spectral distribution of artificial light used in phytotrons has been

criticized because it fails to imitate solar radiation. There are, however,
few data to support this criticism. Elgersma and Meijer (32) in fact present
arguments that sunlight is less effective for plant growth than many ar-
tificial sources currently in use. Nevertheless, some of the plant abnor-
malities encountered in controlled environments have been attributed--
without supporting data--to the spectral composition of the artificial
light, even though as Lang (53) points out it has not been possible to
eliminate the abnormalities by using different combinations of light
sources. Those experienced in the operation of phytotrons know that
light quality is the least likely causal agent of plant growth abnormalities
encountered in controlled-environment rooms. Other environmental fac-
tors and cultural practices are more probable sources of the difficulty.
For example, cauliflower grown in controlled environments often exhibits
a leaf necrosis and a "glassy" curd that rarely are seen under field con-
ditions. Krug et al. (51) have shown that these symptoms result from a
failure to translocate calcium brought about by reduced transpiration.
Normal development was obtained by producing a daily fluctuation in
plant water potential by diurnally cycling relative humidity from 40 to
Table III
Effect o f temperature programming on growth of Bush Blue Lake bean and Samp-
son c u c u m b e r after 15 and 21 days respectively.
Stem length Fresh weight Leaf area

(cm) (g) (cm -~)
Bean Cucumber Bean Cucumber Bean Cucumber
Programmed 7.8b3 27.4b 11.08b 28.67b 280.5b 683. lb

Day/Night' 15.2a 55. la 18.98a 39.72a 567. la 901.8a
Constant 2 18.7a 55.0a 28.84a 39.21a 657.2a 936.8a
' 29.5/23~ the average of the programmed temperatures during the day and night parts
of the diurnal cycle.
2 26.5oc, the average of the 24 hr programmed temperatures.
3 Mean separation within columns by Duncan's Multiple Range Test, 50% level.
90%. Improper selection of temperature can cause nitrogen deficiency

symptoms and leaf edge burn similar to that produced by sodium toxicity
(99), and incompletely calcined clay substrates can lower nutrient solu-
tion pH to intolerable levels (26).
Many ecologists and other field scientists seem to have an unreasoning
dislike for controlled-environment facilities. The reasons for this are un-
clear because few of the objectors have used phytotrons. Some field
scientists seem uncertain that "normal" plants can be grown in controlled
environments and others feel that environmental influences cannot be
understood unless all factors are considered at the same time.
Plants grown in phytotrons are no more abnormal than the investigator
allows them to be. The difference between phytotron and field grown
plants may be no greater than between those grown in the field during a
cooler than normal and an average hot summer. Thus part of the problem
is that we do not know what to call "normal" in the field. Also part of
the problem in developing normal plants is that the concepts of thermo-
and photoperiodism have created what is often an overemphasis on diur-
nal variations. For example, programming temperatures to simulate an
average day during the growing season may seem, without further in-
spection, to be a valid procedure; even though it has little ecological
significance. Unfortunately the diurnal program developed from weather
records frequently results in plants that grow more poorly, or at best no
better, than those at constant temperatures (Table III). It is possible,
however, to produce plants in the phytotron that are phenotypically sim-
ilar to those in natural environments (76). This is done by simulating the
seasonal progression of environmental events and its accuracy is limited
mainly by the ability of the investigator to define the field phenotype.
Certainly plant growth in the field represents a complex of interactive
PHYTOTRONS 481
processes and to some extent the inability to observe such complexes in

their entirety can restrict the degree to which controlled environment
studies can be correlated with field research and ecology. It seems odd,
however, that those who question the validity of controlled environment
work do not also question studies of isolated, rigidly controlled, bio-
chemical reactions. Yet both controlled-environment studies and bio-
chemistry follow the same basic principle of research----entia praeter ne-
cessitatem non sunt multiplicanda. (The number of entities should not be
increased unnecessarily.)
PHYTOTRON RESEARCH
Research in phytotrons is extremely varied. For example, the research

at the SEPEL phytotron has been reported in over 40 different journals
representing such diverse fields as air pollution, agronomy, botany, hor-
ticulture, entomology, nematology, pathology and physiology. Most phy-
totrons can provide lists of publications and occasionally general descrip-
tions of the research are published (6). Climate Lab News keeps the
reader up-to-date on the research at Palmerston North and annual reports
are available from CERES and SEPEL. Summaries of the work in various
phytotrons also appear in the Phytotronic Newsletter (14, 41, 49, 86).
Digests of the research at IVT Wageningen (83) and Taiwan (52) have
been compiled, and Biotronics regularly reports the major research con-
ducted at the Biotron Institute in Japan.
Biological and Technical Service

Phytotron staffs regularly conduct experiments designed to evaluate
and improve experimental methods. For example, techniques are being
evaluated to simplify the maintenance of a species collection. The usual
method for replacement of the seed supply is to place species from di-
verse habitats in a glasshouse and grow them until they flower and set
seeds. Since the environment is rarely optimum, and in fact during part
of the year can inhibit reproduction, the process is inefficient. Optimum
conditions for flowering and seed production can be determined, how-
ever, by using a phytotron to provide a series of temperature and day-
lengths. Seeds of Nicotiana thyrsiflora, for example, are difficult to ob-
tain in a glasshouse because the plants fasciate severely. The fasciated
plants produce few flowers and many of these are abnormal. Standard
phytotron techniques showed clearly that fasciation was a consequence
of high temperatures and long days. Short days and day/night tempera-
tures of 22/18 or 18/14 produced enormous numbers of seeds on normal
inflorescences (28).
Table IV
Effect of high CO2 concentrationson growth and developmentof several species.
Fresh weight (g)

No. of
Dura- open flowers Flowers + Buds Plant
tion 300 1000 300 1000 300 1000
Species (days) ppm I ppm ppm ppm ppm ppm
Vinca rosea 37 0.2 2.4 -- -- 5.9 9.5
Marigold
Brownie Scout 35 0.9 3.2 0.25 7.75 52.17 75.01
King Tut 37 0.4 2.4 0.30 7.10 61.80 74.76
Sparky 32 0.1 1.1 0.10 2.76 66.53 86.47
Petunia
Comanche 35 0 1.3 -- -- 10.56 22.62
Happy Time 35 0 7.0 -- -- 14.09 29.73
Pinto Bean 22 . . . . 21.66 41.26
Tobacco
Coker 319 25 . . . . 7.94 20.69
i ppm = parts per million.
Increasing the amount of carbon dioxide in the atmosphere increases

growth of some plants (Table IV), but seems to be effective for only
about the first six weeks of growth (31, 87). Decreases in chlorophyll and
leaf nitrate and large increases in starch are commonly observed (29).
Since much basic information needs to be learned about the role of high
COz levels on whole plant growth, several phytotrons have developed
techniques for using the facilities in this area of research.
Phytotron staffs continuously evaluate methods of improving plant
growth and development. Watering and nutrient application practices are
periodically reevaluated. The effectiveness of various artificial substrates
has been examined in detail and comparative studies have been made of
plant growth under various combinations of light sources at SEPEL and
Palmerston North.
Phytotron technical staffs are continuously at work improving environ-
ment control capability and developing devices to improve data acquisi-
tion. Temperature and humidity controls are redesigned to improve per-
formance and reliability. Vibration, a characteristic of reach-in chambers
using direct expansion refrigeration systems, has been virtually eliminat-
ed at SEPEL. Automatic water and nutrient solution application systems
PHYTOTRONS 483
have been developed and evaluated and at Palmerston North the staff
has designed an effective root washing facility.
A D V A N T A G E S OF P H Y T O T R O N S
Environmental Measurements
Measurement of environmental parameters has long been a problem in
biology. Although the reasons for it are not always clear, biologists have
reported questionable, if not impossible, values for every environmental
factor. A scientist who would insist on elaborate, fast response laboratory
instrumentation often appears satisfied with measuring temperature and
humidity with a hygrothermograph. Moreover the laboratory equipment
is carefully and frequently calibrated and the hygrothermograph is not.
Field scientists place hygrothermographs and other climate station in-
strumentation in shelters constructed according to international standards
set up by the World Meteorological Organization (103). Yet when these
instruments are used in a glasshouse or controlled-environment chamber,
they are simply placed on the plant bench or on a shelf where they will
not interfere with the investigator. One must conclude ipso facto that
many biologists do not understand that an exposed hygrothermograph
can read 6~ or more higher than the air temperature.
Poor reporting of environmental measurements, therefore, seems due
to inadequate instrumentation, failure to recognize the limitations of the
measuring device, and unfamiliarity with the characteristics of the factor
being measured. For example, the overall accuracy of a ventilated psy-
chrometer is _+2%; if it is well designed (8). Relative humidity is depen-
dent upon temperature and changes about 9%/~ at -17 and 5%/~ at
37~ (84). Thus a relative humidity control of _+5% in a controlled-envi-
ronment room filled with plants that are watered two or more times daily
is extremely good and _+3% is remarkable.
Methods of measurement are often controversial. Illuminance, for ex-
ample, is periodically denounced for having no biological significance.
When accompanied by a description of the light source, however, illu-
minance measurements do provide a means of insuring that the light
conditions can be duplicated. Irradiance or total energy measurement,
considered sine qua non by some researchers, suffers from a confusing
array of units and biological relevance is seriously reduced by failure to
report wavelength limits. Light measurements also suffer from infre-
quently calibrated instruments. These problems were discussed in detail
at a symposium to establish controlled environment guidelines (88).
Phytotrons, as a rule, are equipped with better environmental measur-
ing equipment than can be afforded by the individual plant growth cham-
ber or glasshouse operator. Moreover, the phytotron staff keeps the in-
struments calibrated and experienced phytotronists make valid
measurements.
Efficient Maintenance and Operation

The central administration of a large group of controlled environment
facilities afforded by phytotrons reduces wastage of research space due
to idle periods and excessive down-time for repairs and maintenance.
Since phytotrons have full-time maintenance staffs, scientists can do their
research without also functioning as electricians and mechanics; func-
tions for which they may have little training and less enthusiasm. From
a technical standpoint, phytotrons allow research biologists to collaborate
closely with design personnel to provide the most effective integration of
skills in developing new or specialized facilities. As an example, a sci-
entist recently ordered a simple, walk-in incubator. Space limitations re-
quired the unit to be installed out-of-doors where, although protected
from rain, it will still be subjected to drastic seasonal temperature
changes. Although the specifications were clear and unambiguous, the
lowest bidder failed to meet them. The scientist has now spent many
hours of research time trying to get the equipment operable and the delay
has set back his research program considerably. The real cost of the
equipment thus has increased enormously. Many phytotrons could pro-
vide facilities for this investigator without difficulty or loss of time. If
incubator type equipment was not available, it could be added by per-
sonnel experienced in the problems of environment control in biology.
Moreover, the characteristics of secondary coolant refrigeration systems
would allow the incubators to be operated without significantly increasing
the total power consumption.
Biological Aspects
Several views of the importance of phytotrons to basic and applied
research were discussed earlier (15, 20, 33, 50, 53, 62, 63, 99). Phytotron
research is predominantly disposed to investigations with whole plants
but this does not mean that biochemistry and molecular biology are ig-
nored or that no basic research is done. In fact, a phytotron is ideally
suited for studies of the mechanisms by which differences in metabolic
patterns are influenced by environment. Phytotron research, however,
usually studies biochemical events with regards to whole plant or popu-
lation behavior instead of as isolated systems.
Phytotrons provide a range of controlled conditions that allow com-
prehensive factorial experiments to determine multidimensional interre-
lations of environmental and physiological systems. Thus as Paul
PHYTOTRONS 485
Chouard has pointed out, in a phytotron the gap between physiology and
microclimatology can be narrowed, ecology can become an experimental
science, the genetics of response to climate can be explored and the
biochemical and physiological mechanisms affected by climatic stress can
be identified and understood.
It should be clear that phytotrons are research tools designed to answer
questions about the role of environment in all phases of plant develop-
ment. Whether or not the answers contribute sensible information to
basic and applied science depends, like all research, in asking the right
questions in the right way.
ACKNOWLEDGMENTS
The directors and staffs of the various phytotrons generously supplied

information about their facilities. Practicalities, however, allowed inclu-
sion of only a few of the schematics and photographs supplied by N. de
Bilderling in France, Helmut Krug in Germany, T. Matsui in Japan, Stein
Nilsen in Norway, E. O'Neill in Australia, and Ian Warrington in New
Zealand.
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THE BOTANICAL REVIEW

VOL. 46 OCTOBER-DECEMBER, 1980 No. 4

Abstract ........................................................................................... 447

Phytotrons are the most complex form of controlled-environment fa-

The BotanicalReview 46: 447--489, October-December, 1980.

Los fitotrones son la forma mas completa de "ambiente controlado"

en varias fases de la experimentaci6n ecol6gica. Ya que las plantas pue-

"Environment seems to be the key word in an amazing number

reproducible. Since no amount of statistical manipulation can substitute

Fig. 2. Early, 1947, controlled-environment room equipped with slimline fluorescent

Fig. 3. Controlled-environment room constructed in 1963 at Beltsville using highly load-

A phytotron is often visualized as consisting of a combination of tem-

3,600 L 1,800 ] 3,600

ature and humidity controlled glasshouses and several phytotrons are

advised on the development of new phytotrons at Odessa, Mironovka,

C11 C12 C13

C21 C22 C23

: "-. ,b22 I "- .. ib23

C21/ll~_ / , , _ . , / C22,ft~ 60 F- dt-C23

I., -d~. ', \ ~,, ', " \ ", / / / / / Lip_- 23

a21-'r "ra2 2 "ra2 3

demand. Hot water is produced by solar heating panels located on the

Controlled Environment Chambers

Most phytotrons control air temperature. Radiation effects are elimi-

looked is that their primary application is to provide low relative humid-

The Phytotron Building

co~vl~r,~ ~IIIR T.Ip

General Design Criteria

maturely (25). Moreover, incandescent lamps are inefficient light pro-

Phytotrons may have no permanent research personnel other than the

processes of morphogenesis to biochemical mechanisms and related bio-

A major criticism of early phytotrons was the lack of flexibility in the

Fig. 11. G r o w t h of B u s h Blue L a k e b e a n placed in a 2.44 x 3.66 m controlled-environ-

The spectral distribution of artificial light used in phytotrons has been

Stem length Fresh weight Leaf area

Programmed 7.8b3 27.4b 11.08b 28.67b 280.5b 683. lb

90%. Improper selection of temperature can cause nitrogen deficiency

processes and to some extent the inability to observe such complexes in

Research in phytotrons is extremely varied. For example, the research

Biological and Technical Service

Fresh weight (g)

Vinca rosea 37 0.2 2.4 -- -- 5.9 9.5

Pinto Bean 22 . . . . 21.66 41.26

i ppm = parts per million.

Increasing the amount of carbon dioxide in the atmosphere increases

Efficient Maintenance and Operation

The directors and staffs of the various phytotrons generously supplied

14. Champagnat, P. 1976. Actual research at the CNRS phytotron in Gif-sur-Yvette,

Chamber Environments 20th Internatl. Hort. Cong. Phytotronic Newsletter 19:

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