Journal of Horticultural Science

ISSN: 0022-1589 (Print) (Online) Journal homepage: http://www.tandfonline.com/loi/thsb19

Effects of humidity on the growth and yield of

glasshouse tomatoes

Rachel Holder & K. E. Cockshull

To cite this article: Rachel Holder & K. E. Cockshull (1990) Effects of humidity on the
growth and yield of glasshouse tomatoes, Journal of Horticultural Science, 65:1, 31-39, DOI:
10.1080/00221589.1990.11516025

To link to this article: http://dx.doi.org/10.1080/00221589.1990.11516025

Published online: 27 Nov 2015.

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 Journal of Horticultural Science (1990) 65 (1) 31-39

Effects of humidity on the growth and yield of glasshouse

tomatoes
By RACHEL HOLDER
Efford Experimental Horticulture Station, Lymington, Hampshire S041 OLZ UK*
and K. E. COCKSHULL
AFRC Institute of Horticultural Research, Littlehampton, West Sussex BN17 6LP, UK

SUMMARY
Four discrete humidity treatments (0.1, 0.2, 0.4 and 0.8 kPa vapour pressure deficit) were
maintained continuously for 28 days from mid-January and their effect on the growth and
yield of a long-season tomato crop was investigated. The average vapour pressure deficits
achieved over the 28 day period were0.15, 0.25, 0.43 and 0.65 kPa for the four treatments
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respectively. Temperature differences between the treatments were less than 0.2°C. The
rate of plant development was unaffected by humidity. Significant reductions in leaf area
at high humidities were associated with low calcium concentrations in the leaf laminae
and calcium deficiency symptoms. There were no calcium deficiency symptoms in the
fruit. The trusses associated with the smaller leaves resulting from high humidity, pro-
duced smaller fruit and thereby a lower yield compared with the low humidity treatments.
Fruit yields produced before and after these trusses were picked, were unaffected by the
humidity treatments. High humidity also reduced fruit quality. It was concluded that the
cost of reducing humidity to vapour pressure deficits greater than 0.3 kPa was likely to
exceed any economic gain, because the yield response to lower humidities was very small.

THE influence of humidity on the growth and avoid these disorders, costly humidity control is
r- yield of protected crops is poorly understood. practised in commercial production. However,
This deficiency in our knowledge has been cost-effective humditity control cannot be
identified as a constraint on optimization of realized until the full effect of humidity on crop
crop production in insulated greenhouses productivity is known.
(Cockshull, 1985). There have been few studies High humidities can be expected to suppress
of humidity effects on crop productivity (e.g. transpiration and thereby reduce the uptake of
Bakker, Welles and Uffelen, 1987) and in a nutrients in general (Adams, 1980) and the
review of the research to date, Grange and transport of calcium in particular (Stebbins and
Hand (1987) · concluded that humidities Dewey, 1972). High humidities are also associ-
between 1.0 and 0.2 kPa vapour pressure defi- ated with leaf-scorch symptoms and reduced
cit (vpd) had little effect on the physiology and effective and actual leaf areas (Bakker, 1984;
development of horticultural crops. However, Starkey, 1985) which may lead to smaller fruit
vapour pressure deficits smaller than 0.2 kPa size and yield. Bakker (1984) has demonstrated
can be produced for extended periods in mod- that these leaf symptoms are associated with
ern glasshouses which are well-sealed and localized calcium deficiency.
incorporate energy-saving measures, such as Glasshouse experimentation with accurate
thermal screens, fixed screens, or double glaz- humidity control, independent of associated
ing (e.g. Starkey, 1985) Such high humidities changes in temperature or other environmental
can promote the incidence of calcium-related, factors, is difficult and may be the reason why
physiological disorders (Bakker, 1984). To there have been relatively few studies of
*Present address: Van Heyningen Bros. Ltd., Toddmgton humidity and crop productivity. These prob-
Lane, Littlehampton, West Sussex, BN17 7PP lems have largely been overcome in a purpose-
 32 Humidity and glasshouse tomatoes

built, multi-compartment glasshouse sited at tration was measured with a Siemens analyser
Efford Experimental Horticulture Station, and and pure C0 2 used to maintain 1000 vpm from
the first experiment to use this facility is dawn to dusk in all compartments, regardless of
described here. ventilator position, until 12 February 1987.
Thereafter, the concentration was maintained
MATERIALS AND METHODS
at 1000 vpm until the ventilators were more
The Venlo-type glasshouse unit used in this
than 5% open, when it was controlled at
study comprised 16 compartments arranged to
350vpm.
form a four by four array, surrounded by a
Four different humidity treatments (0.1, 0.2,
heated corridor with double-glazed external
0.4 and 0.8 kPa vpd) were maintained continu-
walls (Figure 1). All compartment walls were
ously for 28 days from 15 January to 12
internal, therefore, with only a small temper-
February 1987. Each treatment was replicated
ature gradient across them, features which
in four compartments randomized within a
make the facility ideal for humidity studies.
Latin Square design. The 0.8 kPa vpd treat-
Each compartment (8.0 m x 9.6 m) was
ment was the control. After the period of treat-
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equipped with computer control (Brink~~n,

ment ended, the humidity in all compartments
Bucom) of air temperature and humidity.
was maintained at a vpd of 0.3 kPa or more. At
Humidity was lowered by applying minimum
the start of treatment, one half of the flowers of
ventilation and additional pipe heat, whilst
the third truss had reached anthesis, while by
atomization of softened water (fogging) was
the end of treatment, the first flowers of the
used to increase it. This tended to reduce air
seventh truss had reached anthesis.
temperature, and so temperature set points
Tomato seeds (Lycopersicon esculentum Mill
were adjusted at intervals to maintain the ~arne
cv. 'Counter') were sown on 24 October 1986
average day, night and 24 h temperature m. all
and propagated on rockwool using standard
compartments. Humidity was mon~tored usi?g
techniques (Starkey, 1985). The young plants
aspirated wet- and dry-bulb platmum resis-
were transferred to their final cropping
tance thermometers. Their accuracy was reg-
locations on 1 December 1986 and the pro-
ularly checked using an Assman psychrometer
pagation cubes were brought into contact with
and found to be within ± 0.1 oc of the psychr-
ometer reading. Carbon dioxide (C0 2) concen-

1 2 3 4
N

5 6 7 8

48m 1 9.6m

9 10 11 12

13 14 15 16

t--1,.------ 40m------- Sm -------1.,~1

FIG. la
The arrangement of compartments within the glasshouse FIG. lb
unit. The arrangement of plant rows within a compartment.
 RAcHEL HoLDER and K. E. CocKSHULL
the rockwool slabs on 17 December 1986.
Initially, the volume of nutrient solution
applied to the slabs was 50% more than that
required to offset the uptake and loss of water.
This was necessary in order accurately to main-
tain the desired calcium concentrations. Later,
irrigation frequency was directly linked to solar
radiation receipt and the total volume applied
each day was based on estimates of potential
evapotranspiration using the standard Penman
equation. This was adjusted, as necessary, to
provide 30% excess solution, measured as 'run
off from the rockwool slabs.
Within each compartment, the plants were
arranged in four double rows, aligned S-N with
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17 plants per row to give a density of2.65 plants
m·2 • Single rows of plants were arranged along
the length of the east and west walls of each
compartment to act as guards (Figure 1). All
plants were grown to a height of about 3.4 m
and then layered. Whitefly were controlled by
introducing the parasitic wasp Encarsia for-
mosa and, every two weeks, sprays of fungi-
cides were routinely applied to all plants using
'Elvaron', 'Rovral', and chlorothalonil in
rotation.
Each double row within a compartment
received a different nutritional sub-treatment,
applied as one of four nutrient solutions con-
taining either 150 or 300 mg Ca 1" 1 (to give
approximately 200 or 400 mg Ca 1" 1 in the rock-
wool slabs) combined with one of two conduct-
ivities (to .give either 5 or 7 mS cm· 1 in the
rockwool slabs) in a two by two factorial
design. The conductivity sub-treatments were
maintained from the time that contact was
established between the propagation cube and
the slab until two weeks before fruit were
picked. Conductivity levels were then lowered
gradually to 3 and 5 mS cm· 1 respectively, four
weeks after the onset of picking. The concen-
trations of all nutrients other than calcium and
sulphate were kept the same in all sub-treat-
ments by raising the conductivity of the
solutions to the desired levels with the addition
of sodium chloride (Adams and Bailey-
Anguish 1988). The nutritional aspects of this
study will be reported elsewhere and so, the
data presented here are the means of four
sub-treatments.
Calcium deficiency of the tomato leaves was
assessed visually on 24 February 1987, as
33
described by Holder and Cockshull (1988), and
samples of young leaves (fifth leaf below the
head of the plant) were taken at intervals for
routine tissue analysis. Leaf area was assessed
indirectly by measurement of maximum leaf
length and width for the leaf immediately
below trusses one to 14 inclusive. This assess-
ment was carried out on the 5.0 mS cm· 1 :
150 mg Ca 1" 1 sub-treatment.
Fruits were picked three times a week from
each nutritional sub-treatment and the weight
in each Class (I, II and waste) and Class I size
grade (35-41 mm, 42-47 mm, 48-57 mm and
58-67 mm in diameter) recorded. The inci-
dence of fruit disorders was estimated on fruit
from one half of the plots (the northern eight
compartments) on 1.4.87. The percentage
compression of fruit picked on 25.3.87 was
determined after six days in a shelf-life room
(20°C, 60% rh). The test used samples of 20
fruit (47-57 mm diam.) from each sub-treat-
ment and they were tested on a standard 'firm-
ness meter' (Hobson and Ambler;1988).
RESULTS
Aerial environment
Accurate control of humidity was not always
obtained but, nevertheless, a wide range of dis-
crete humidity regimes were produced in the
compartments (Figure 2). The values for aver-
age temperature, humidity and calculated solar
radiation integral at crop height in each envi-
ronment (Table I) demonstrate that humidity
control was largely independent of other
factors.
Calcium deficiency
Calcium deficiency symptoms were first seen
14 days after the start of the humidity treat-
ments and were restricted to the 0.1 and
0.2 kPa vpd treatments. The initial symptoms
appeared as a lime-green discoloration of the
terminal leaflet margins of the rapidly expand-
ing leaves and an associated reduction in their
area. Affected leaves expanded slowly and
their leaflet margins subsequently became chlo-
rotic and bleached. In severe cases the disorder
developed interveinally towards the midrib and
the leaflet became cupped. Calcium deficiency
symptoms could develop, however, at any stage
of leaf expansion but their spread halted almost
immediately after the high humidity treatments
 34 Humidity and glasshouse tomatoes

.···...·...·. ·.. ........ .. ····...... .. ... ······.......... . ...... .····.

~· "•. •''•

0.2

ca
a.
~

.ti
u
'i 0.4
~

f
:I

..
::1Ill
Q. 0.6
:I
Downloaded by [UNSW Library] at 10:29 26 October 2017

0
Q.
ca
>

0.00 12.00 0.00

Time (25.1.87)

FIG. 2
Records of humidity (kPa, vapour pressure deficit) from the four treat-
ments (compartments 1 to 4) over one pcnod of 24h.

were stopped. Consequently, the deficiency Rate of plant development

symptoms were restricted to leaves serving Neither the rate of plant development, as
trusses four to seven. Symptom expression was measured by the rate of truss production, nor
most severe at the highest humidity and analy- plant height were affected by the humidity
sis of the nutrient content of leaf laminae con- treatments. The rate of truss production
firmed that the disorder was associated with a between 16 December and 13 February aver-
deficiency of calcium (Figure 3). Samples from aged 0.78 (± 0.02) trusses per week, while
young leaves, taken after the humidity treat- plant height on 13 February was 2.0
ments had ended, showed that, although the (± 0.01) m.
percentage calcium content was no longer
correlated with treatment, the amount of cal- Leaf growth
cium per leaf was still at a low level in the high The growth of leaves serving trusses four to
humidity treatment (Adams, 1988). ten (inclusive) was significantly reduced by high
TABLE I
Average aerial humidity, temperature and calculated solar radiatwn integral at crop he1ght m three different periods. Period
1=17.12.86-14.187, 2=15.1-/2.2.87; 3=13.2-31.3.87
Humidity (kPa vpd) Temperature (0 C) Radiation mtegral (MJ m- 2 d-1)
Treatment
(kPa vpd) 2 3 2 3 2 3
0.1 0.39 0.15 0.45 17.6 18 1 18.5 !.55 1.80 4 85
0.2 0.41 0 25 0.45 17.6 18.1 18.4 1.55 1.83 4.85
0.4 0.41 0.43 0.46 17.6 18.1 18.4 1.55 1.89 4.85
0.8 0.39 0.64 0.46 17.6 18.1 18.4 1.55 1.91 4.85
 RACHEL HoLDER and K. E. CocKSHULL 35

...
Ul.c
Qll:ll
>.-
CIIQI
..!!~
0~
.. "C
c~
Qll
.. 1:11
SE
1.1::1
E1!
:lea
·u ~,~
-~:~~

~E

0 0.2 0.4 0.6

Average vapour pressure deficit achieved, kPa
FIG. 3
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The relationship between humidity, calciUm deficiency symptoms in

the leaves of tomato on 24 2 87 ( • ), and their calcium content ( o)
SED for deficiency symptoms=O 311 (3 d.f.).

humidity (Table II). The pattern of leaf size fruit size. Indeed, the total weight of fruit
relative to that in the 0.8 kPa vpd treatment is picked in the two smallest size grades between
shown in Figure 4. The smallest leaves were 21 March and 22 May was either unaffected or
below trusses five and six. These leaves were increased by high humidity, whereas that in the
expanding at the time the humidity treatments two largest size grades was significantly
were applied. On average, the flowers of truss reduced (Table IV).
four had reached anthesis one week after the
Fruit quality
start of treatment, while those on truss seven
High humidity did significantly affect the
were at anthesis by the end of the four-week
yield of Class I fruit prior to late March, when
treatment period. Thereafter, the differences in
total yields were unaffected, and during the late
leaf size became less marked and there were no
March to late May period when both total and
consistent residual effects of humidity on leaf
Class I yields were lower from the plants that
size between trusses 11 and 14.
had had the higher humidity treatments (Table
Fruit production V). High humidity did not affect the incidence
Picking of fruit began on 16 February, just as of fruit disorders (data not presented) and
the humidity treatments ended. Yields for the TABLE II
first five weeks of picking were unaffected by The effect of humidtty treatments on the product of leaf length
the humidity treatments but the weight of fruit (em) X leaf wzdth (em) for the leaf zmmedwtely below trusses
I to 14 (trusses 1--{5, assessed 3.3.87; trusses 7-14, assessed
picked over the period from 23 March to 22 13.4.87)
May was significantly reduced by high humidity
Humidity treatment
(Table III). This period coincided with the pick- (kPa, vpd)
ing of fruit from trusses four to ten which were Leaf SED
the trusses associated with smaller leaves position 0.1 0.2 0.4 0.8 (6 df) Sigmficance
resulting from the high humidity treatments. 1 1899 1965 1886 1903 87.6 n.s.
Thereafter, the humidity treatments had no 2 2307 2333 2213 2300 97.7 n.s.
3 2119 2129 2074 2183 128 8 n.s.
significant effects on either yield (Table III) or 4 1534 1708 1887 2040 67.9
leaf size (Table II). In general, the effects of 5 1159 . 1364 1817 2022 78.6
increasing humidity on yield and on leaf size 6 1034 1294 1666 1955 58.3
7 1372 1712 1970 2125 83.5 •••
were similar.
The main components of yield are fruit
8
9
10
1381 1656
1638 1892
1876 1997
1871
1952
2052
2038
100.6
104.1
..
.
number and fruit size. The humidity treatments 2090 2265 72.4
11 2234 2192 2302 2265 115.0 n.s.
had no effect on the numbers of fruit set on 12 2453 2193 2295 2382 55.2
trusses four to ten (9.19 ± 0.22 flowers per 13 2278 2032 2114 2248 123.6 n.s
14 1964 1777 1898 2010 118.4 n.s.
truss) and so their main effect was probably on
 36 Humidity and glasshouse tomatoes
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Truss number

4 FIG.
The effects of humidity on the s1ze (length x breadth) of leaves below successive fruit trusses,
relative to that in the low humid1ty treatment (0.8 kPa, vpd). 0.4 kPa (o); 0.2 kPa (A); 0.1 kPa
(•).

although the firmness of fruits that were plays a major role in determining the early yield
developing while the humidity treatments were of tomato crops although total yields are less
given were, on average, slightly softer, the affected (Slack and Calvert, 1978). It is essen-
effects were not significant nor consistent tial, therefore, that temperature differences
within sub-treatments (data not presented). between humidity treatments are minimized if
unequivocal comparisons of growth and yield
Diseases are to be made in different humidities (e.g.
Despite the high atmospheric humidities that Bakker et al., 1987). The lack of treatment
were used, no fungal infections were observed. effects on the rate of truss production confirm
that equality of temperature integrals was
DISCUSSION achieved (Table I). The major remaining prob-
Temperature integral from time of flowering lem is the possible confounding of light integral
TABLE III TABLE IV
The effect of humtdlly treatments on total fruit yield m three The effect of humidity on the Class I fruit size distribution by
periods (1=12.2-20.3.87; 2=21.3-22.5.87; 3=23.5- weight between 21.3 and 22.5.87
30.9.87)
Weight (kg m -z) in each diameter size grade
Yield (kg m- 2) during period (mm)
Treatment Treatment
(kPa vpd) 2 3 (kPa vpd) 35-42 42-47 47-57 57-67
0.1 1.35 10.1 33.8 0.1 0.20 2.0 6.4 0.11
0.2 1.28 10.7 33.8 0.2 0.17 1.8 7.1 0.17
0.4 1.31 11.1 33.8 0.4 0.14 1.9 7.4 0.22
0.8 1.36 11.4 33.9
SED
Significance
0.057 (6 d. f.)
n.s.
0.19 (6 d.f.)
**
0.19 (3d. f.)
n.s.
0.8
SED (6 d. f.)
Significance
0.12
0.012
1.8
0.072
n.s. ...
7.7
0.25
0.43
0.073
 RACHEL HoLDER and K. E. CocKSHULL
TABLE V
The effect of humidity treatments on Class I fruit yield in three
periods (1=12.2-20.3.87; 2=21.3-22.5.87; 3=23.5-
30.9.87)
Class I yield (kg m- 2) during period
Treatment
(kPa vpd) 2 3
0.1 0.77 8.7 29.9
0.2 0.77 9.3 30.0
0.4 0.81 9.7 30.0
0.8 0.88 10.0 30.1
SED (6 d.f.) 0.022 0.255 0.36
Significance n.s.
with humidity treatment. In the present study,
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greater condensation on the internal surface of
the glasshouse roof in the most humid com-
partments and the presence of small (5-10 J.tm
diameter) water droplets which were injected
into the aerial environment (to raise the humid-
ity) as frequently as every four minutes, would
both have reduced light receipt by the crop in
the high humidity treatments. Long-term
measurements of light receipt in the different
humidity treatments were not available, but
some short-term measurements suggested that
a maximum of 6% light loss occurred in the
0.1 kPa vpd treatment (Table I). Data of Cock-
shun and Graves (Cockshull, 1988a) indicate
that a light integral loss of this magnitude at this
time of year and prior to picking fruit would
have minimal effect on yield (to a maximum of
0.1 kg m·2). For this reason no attempt has been
made to adjust the yield data.
The dramatic effects of humidity on the
expression of calcium deficiency symptoms
have been unambiguously demonstrated for
the first time. It had previously been shown that
the calcium content of leaves was reduced in
tomato plants grown in controlled environment
cabinets at high humidity (Gislerod, Salmer-
Olsen and Mortensen, 1987; Adams, 1988;
Burrage, 1988) and symptoms of calcium defi-
ciency had been induced in tomato crops grown
in high humidity environments (e.g. Buitelaar,
1985; Starkey, 1985). These symptoms together
with associated reductions in leaf area were
attributed to the effects of elevated humidity
levels, although air temperatures had also dif-
fered in the latter experiments.
A remarkable result from this study is the
difference in the effect of humidity on leaf size
37
and yield. Leaf area was reduced by up to 50%
at the highest humidity treatment compared
with the lowest, whereas yield was never
affected by more than 31%. Previous humidity
studies on tomato crops using a smaller range of
humidities and a lower maximum humidity,
although over a longer period, have reported
both greater and smaller yield reductions asso-
ciated with high humidity (Buitelaar, 1985, and
Winspear, et al., 1970, respectively). In the
present experiments, the trusses that were most
affected were those associated with smaller
leaves resulting from the high humidity. This
would be expected if the assimilate supply to a
truss comes principally from leaves in its
immediate vicinity (Ho and Hewitt, 1986).
The response of tomato to humidity was
markedly different from that of cucumber and
sweet pepper as reported by Bakker (1988).
Cucumber produced larger leaves at higher
humidities up to 0.4 kPa vpd; the highest
humidity investigated by Bakker eta/., (1987).
Furthermore, observations in the present study
showed that a reduction in leaf area of toma-
toes occurred before visible symptoms of cal-
cium deficiency. Indeed, smaller leaves may be
produced in treatments such as 0.4 kPa vpd, in
which almost no deficiency symptoms
appeared. This contrasts with the growth of
cucumber leaves which was enhanced at high
humidity despite the presence of calcium defi-
ciency symptoms (Bakker eta/., 1987). Burrage
(1988) demonstrated that high humidity also
reduced the leaf area and dry weight gain of
young tomato plants. The uptake of various
ions was also reduced but in proportion to the
reduction in dry weight.
There was no evidence that humidity
affected either pollination or fruit growth
directly or the incidence of fungal disease, and
so the yield differences could be attributed to
differences in fruit size; a finding similar to that
reported by Lipton (1970) and Starkey (1985).
It is of interest that others (e.g. Wins pear eta/.,
1970; see also Grange and Hand, 1987) have
reported a greater incidence of fungal disease
at high humidity. A number of factors probably
contributed to its absence in the present experi-
ments. First, as water was introduced into the
atmosphere in the form of a 'fog' of fine drop-
lets and as the humidity was controlled at less
than saturation, films of moisture were less
 38 Humidity and glasshouse tomatoes

likely to form on the exposed surfaces of the
plants. Furthermore, the routine application of
fungicide sprays and the use of a cultivar with
genetic resistance to certain diseases also
helped.
There are at least two possible reasons why
the yield depression at high humidities was less
than the associated loss of leaf size. The solar
radiation that was not intercepted by the
smaller leaves formed at high humidities, may
have been intercepted by older, lower leaves
which then provided a limited supply of assim-
ilates to higher trusses, as had been inferred
from leaf removal experiments (e.g. Slack,
1986). Secondly, the leaf area:yield response
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to be economic at vapour pressure deficits
greater than 0.3 kPa, because the yield
response at lower humidities is very small. A
model of the transpiration processes suggests
that the minimum acceptable vapour pressure
deficit will depend on the solar radiation avail-
able (Aikman and Houter, 1990). However, if
the goal of humidity studies is to seek the econ-
omic optimization of crop production, then
further research is needed on the short-term
effects of day and night humidity levels and
their interactions with solar radiation on trans-
piration rate. Indeed, as transpiration is the
plant process with which humidity experimen-
tation is primarily concerned, it should become

may have been less than unity under the experi-
mental conditions with a large winter/spring
canopy and relatively low light integrals.
Unfortunately, no light interception data are
available.
It has been reported that fruit grown at high
humidity are less firm (Buitelaar, 1985) but this
was not confirmed in the present studies.
Unlike several previous studies (e.g. Lipton,
1970), there was also no evidence that fine net
cracking (russetting) was more pronounced at
higher humidities.
From this first study of the relationship
between discrete humidity levels and the
growth and yield of a tomato crop it is con-
cluded that costly humidity control is unlikely
the focus of future experimentation and could,
eventually, become a controlled variable in
commercial tomato production (Stanghellini,
1987, 1988; Cockshull, 1988b).
The authors are grateful to Mr. R. Edmond-
son for the experimental design and the analy-
sis of data, and to Mr. R. F. Clements, Director
of Efford Experimental Horticulture Station
and many other colleagues within the Agri-
cultural Development and Advisory Service
and the Institute of Horticultural Research,
Littlehampton for advice, assistance and infor-
mation. This work forms part of the ADAS/
AFRC Collaborative Project on Energy Saving
in Greenhouses.

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 RACHEL HoLDER and K. E. CocKSHULL 39

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(Accepted 8 Sept~mber 1989)