/. Amer. Soc. Hort. Sci. 101(1):70—73. 1976.

Growth of Lettuce Under Controlled Humidity Levels1

T. W. Tibbitts and G. Bottenberg2
Department o f Horticulture, University o f Wisconsin, Madison, W I53706

A dditional index words. Lactuca sativa, stom ata, stom atal resistance

A bstract. The influence of atmospheric moisture levels of 85% and 50% relative humidity (RH) (3.5 and 11.6
mb saturation vapor deficit respectively) at 20°C for 16:8 long day (LD) cycle on butterhead lettuce, {Lactuca
sativa L. cv. Meikoningen), during a growth period of 4 weeks from seeding, was studied in organic soil and liquid
cultures. Significantly faster growth rates were evident on plants developing under 85% RH than under 50% in
both soil and liquid culture with the largest increases occurring with plants grown in soil culture. The higher
humidity level increased leaf number 15%, leaf size 30%, dry weights 62%, and leaf water contents from 93% to
94%. The differences in leaf number and dry weight equalled about 2 days growth difference at 4 weeks after
seeding. The density of stomata was greater on plants grown under 50% RH but the total number of stomata per
leaf under the 2 humidity levels was the same. The leaf resistance was significantly higher on plants grown under
50% than under 85% RH. The principle significance of high humidity level during growth of lettuce is the produc­
tion of larger marketable heads with a higher water content in a slighly reduced period of time.

Air humidity level has been shown to effect plant growth was 10—17 m min- 1 at the top of the plant canopy as
through its influence on tissue moisture stress. Moisture stress measured with a hot wire anemometer. Fresh make-up air was
increases in proportion to the saturation pressure deficit of the provided and CO2 was monitored at intervals to insure that it
air, and many species exhibit reduced growth under increasing did not go below 300 ppm during the light period.
air moisture deficits. Differences of 5—10 mb (20—40% RH ‘Meikoningen’ lettuce, a butterhead type, was grown in
difference at 20°C) greatly influence the growth of beets, liquid culture and in organic soil, in separate studies. For liquid
peppers, tomatoes, corn, cotton and apples (1, 2, 3, 5, 12), culture, plants were seeded at 20°C on slanted sheets of
and to a lesser degree the growth of ageratum, petunia, germination paper. After 4 days, the seedlings were transplanted
marigold, radish, wheat, soybeans and sugar beets (1, 3, 6, 11). into 1-liter containers of nutrient solution continuously aerated
Snapbeans, barley, peas and cacao have shown no significant and continuously supplied with 200 ml per minute of solution
effects from increased humidity levels (1 ,7 , 8, 9). from a 240-liter tank. Excess solution drained back into the
Increased humidity levels have produced plants with tank for recirculation. The 240-liter supply provided adequate
increased vegetative growth resulting in increased flowering and nutrient for the 10 plants in each chamber for the total period
seed yields (1). With certain species, such as cotton, highest of an experiment as determined by solution conductivity
humidity levels interfered with fertilization and fruit develop­ measurements. The solution contained concentrations of
ment, reducing the seed yield (3). Increased growth from nutrients as follows:
increased humidity may result either from reduced stomatal Meq per liter Parts per million
resistance and thus increase CO2 uptake (10) or from increased Na 0.5 Fe 2.5
cell enlargement which provides a greater amount of leaf area K 3.0 B 0.25
for light absorption (3). Mg 2.0 Mn 0.25
These studies were undertaken to determine if atmospheric Ca 5.0 Zn 0.025
moisture will alter growth rates of lettuce when grown either in no3 7.5 Cu 0.01
soil or liquid media and to provide insight into the physiological SO4 2.0 Mo 0.005
basis for growth responses. H2P 04 0.5
experiments in organic soil, native peat soil from a
Materials and Methods Wisconsin marsh with a pH of about 6.0 was utilized. It was
Studies were conducted in M-3 Environmental Growth moistened with nutrient solution to a maximum moisture level
Chambers with plexiglass barriers having 80% input wattage of without puddling. The nutrient concentrations were increased
cool-white 1500 MA fluorescent lamps and 20% input wattage so that at maximum moisture level the soil water contained 8 X
of incandescent lamps that provided a spectral energy distribu­
tion of visible and infra-red radiation as plotted (Fig. 1). The
light intensity at the top of the containers was maintained at
2000 ft-c (21.5 klx) ± 5% as measured with a Weston Model 756
meter. Light duration was 16 hr with an abrupt light-dark
change. Air temperature as measured with thermocouples at the
top of the containers was maintained at 20 ± 1°C during both
light and dark periods to provide a constant saturation deficit
during both periods. Relative humidity, as measured with wet
and dry bulb psychrometers, was maintained in one chamber at
85 ± 2% (3.5 mb of saturation deficit) and in a second chamber
at 50 ± 2% (11.6 mb of saturation deficit) during both the light
and dark periods. The air velocity down through the containers

iReceived for publication April 14, 1975. Research supported by the

College of Agricultural and Life Sciences, University of Wisconsin, Mad­
ison. Fig. 1. Spectral energy distribution of radiation at the leaf surfaces of the
^Present address: Agricultural Dept. Illinois Central College, East Peoria. plants.

70 J. Amer. Soc. Hort. Sci. 101 (1 ):70—73. 1976.

 20 L2

Fig. 3. Enlargement of the fifth leaf of lettuce plants developing under

Fig. 2. Number of leaves developing on lettuce plants for 4 week period 85% and 50% relative humidity when grown in soil culture.
after seeding when grown under 85% and 50% relative humidity in
soil culture.
and attained a larger size. The 5th leaves of plants grown in soil
P and 4 X of the other nutrients compared to the liquid culture. under both humidity levels had passed through the logarithmic
The soil was placed in 3.8—liter metal containers and main­ growth phase by the 25th day after seeding (Fig. 3).
tained within 5% of this maximum moisture level by weighing For plants maintained in liquid culture, the 2nd and 5th
the containers daily and adding the required amount of distilled leaves exhibited approx 15% greater length and width under
water. One plant was grown in each container. 85% RH compared to those under 50% RH. These were signifi­
Duplicate experiments were run in soil culture and in liquid cant differences (Table 1). Thus the total area of individual
culture. The high and low level humidity conditions were inter­ leaves on plants growing under the higher humidity was approx
changed between the 2 chambers in the second study with each 30% greater than for plants under the lower humidity. The 2nd
type of culture. leaf ceased growth completely by 26 days after seeding, while
Growth of lettuce plants was monitored daily or at 3 day the 5th leaf was still showing slow enlargement at this time. The
intervals by counting numbers of leaves and measuring the increase in leaf width of plants grown under the higher humidity
length and width of the second and fifth leaves. Data from 4 or level was slightly greater than the increase in leaf length, thus
more plants was obtained for each treatment in each the leaf length/width ratio of plants grown under high humidity
experiment. Growth rates were compared by calculating mean was slightly but not significantly less than that under low
relative growth rates (RGR). RGR = loge W2 — loge W^/t2—ti humidity.
where W2, is the dry wt at the initial date (tj) and W2 is the dry Fresh and dry wt. The fresh and dry wt of lettuce plants
wt at the final date (t2). grown under 85% RH were significantly greater than those
plants grown under 50% RH. Four weeks after seeding, the
Results fresh wt were 60—100% greater and the dry wt 45—70% greater
The level of humidity under which lettuce plants were grown for plants developing under the higher humidity in both soil and
had a significant effect upon the development of plants grown liquid culture (Table 2). Fresh wt increases with the higher
in either soil or liquid culture. The increase of RH from 50% to humidity were greater than the dry wt increases because the
85% at 20°C (8.1 mb difference in vapor pressure) resulted in amount of water in the tissue of plants developing under the
plant growth with increased leaf number, leaf size, and higher humidity was approx 1% greater (Table 2).
harvested wt. Dry matter accumulated rapidly in lettuce plants during the
Leaf number. The number of leaves on a plant was consis­ 3rd and 4th weeks after seeding and the more rapid rate
tently greater at the higher humidity level in both liquid and soil occurred under the higher humidity level (Table 3). Calculation
cultures. At 25 days the rate of leaf production was approx 1.5 of relative growth rates for g of dry wt per g of dry wt per day
leaves per day at 85% RH and 1.3 at 50% RH (Fig. 2). The over the entire growth period showed a 31% increase per day for
difference was slightly greater in soil than in liquid culture, indi­ plants at 85% RH and 28% increase per day for plants at 50%
cating that the soil was likely imparting a moisture stress upon RH. Therefore, the plants grown at high RH were increasing in
the plants that accentuated the atmospheric moisture stress dry wt at a rate about 11% greater than that in plants growing at
effects. The additive effect of soil moisture stress and atmos­ low humidity. It can also be determined at the relative growth
pheric moisture stress on root crops was demonstrated by Hoff­ rate of 1.31, that the difference in growth between the lower
man and Rawlins (2). humidity plants and the higher humidity plants was equal to
Leaf enlargement. The enlargement of leaves was consistently 2—3 days growing time at 4 weeks after seeding. However, if
greater under high humidity. The leaves grew at a faster rate plants are grown under low humidity to the same dry wt as
plants grown under higher humidity, they will always be
Table 1. Length and width of leaves of lettuce plants grown in liquid smaller, for the leaves will not reach the same size as those
culture for 26 days under 85% and 50% relative humidity. developing under the higher humidity.
Stomatal density. There were 20% fewer stomata cm_2 on
Length of leaf Width of leaf Length/width both the upper and lower surfaces of plants grown under higher
Leaf position (cm) (cm) ratio humidity (Table 4). Stomatal density determinations were ob­
from base 85% RH 50% RH 85% RH 50% RH 85% RH 50% RH tained for leaves 4, 6 and 7 (num bered from the base) and all
showed similar differences between high and low humidity
2 6.35 5.60* 4.50 3.85** 1.41 1.45ns
5 10.70 9.20*
(Table 4). The difference in stomatal density with the 2
8.85 7.45** 1.20 1.23ns humidity levels is in contrast to the similar stomatal densities
* and ** Significantly greater at the 5% level and 1% level respectively found by Slavik (11) with maize grown at 40% and 85% relative
from values at 85% RH. humidities.

J. Amer. Soc. Hort. Sci. 101(1):70—73. 1976. 71

 Table 2. Harvested wt and % water of lettuce plants grown in soil and liquid culture under 85% and 50% relative humidity
from seeding for 4 weeks.
Relative Fresh wt (g) Dry wt (g) % water
humidity
level Soil Liquid culture Soil Liquid culture Soil Liquid culture
(%) culture Expt. 1 Expt. 2 culture Expt. 1 Expt. 2 culture Expt. 1 Expt. 2

85 25.6 19.07 22.81 1.40 1.26 1.33 94.5 93.4 94.2

50 12.7** 9.78** 14.34** 0.82** 0.75** 0.91** 93.4ns 92,3** 93 7**
% increase
with higher
RH 102 95 59 71 68 46 1.1 1.1 .5

**Significantly less at the 1% level than respective values at 85% RH.

Table 3. Dry wt increase of lettuce plants grown under 85% and 50% greater than on leaf 4. This is likely associated with the fact that
relative humidity in liquid culture. leaves 6 and 7 attained a larger size at maturity.
Stomatal resistance. The stomatal resistance of plants
Days after Relative humidity level growing under 85% RH was consistently less than that of plants
seeding 85% 50% growing under 50% RH. The resistance of the lower surface of
all leaves was consistently less than that of the upper surface
(g per plant) (Table 6). The resistance increased with increasing maturity of
15 .044 .039ns the leaves 4, 6 and 7 under 85% RH level but not under 50% RH
21 .309 249**
26 1.292
levels.
.826**
(g per g o f dry wt)
The increased resistance of leaves on plants growing under
Relative growth rate the lower RH indicates less stomatal opening. The fact that
per day for period stomatal density was greater on plants grown under 50% RH,
from 15 to 26 days 1.31 1.28 and a greater number of stomata were present within the area
enclosed by the porometer, indicates an even greater closure of
** Significantly less at the 1% level of significance than respective values
at 85% RH.

Table 4. Number of stomata per cm 2 of leaf surface on lettuce plants developing under 85% and 50% RH.

Leaf numbered from base

Relative _________ 1_______ ________ £________ ________ 1________ _______ Av§z
humidity Upper Lower Upper Lower Upper Lower Upper Lower
level surface surface surface surface surface surface surface surface

85% 6851 5175 6633 6181 7080 6834 6854 6064

50% 8973** 7076ns 7706ns 7471ns 9447ns 8057ns 8409** 7534**

zAverages of 10 counts of .15mm2 per plant from each of 4 plants except italicized values are averages of 3 plants.
*, ** significantly greater at the 5% level (*) and 1% (**) level respectively from values at 85% RH.

Table 5. Total number of stomata on the fourth leaf of lettuce growing Table 6. Stomatal resistance (s-cm 'l) of leaves of lettuce grown under
under 85% and 50% relative humidity. 85% and 50% relative humidity.

Relative humidity level Upper leaf surface Lower leaf surface

Variable 85% 50% Leaf number RH level RH level
from base 85% 50% 85% 50%
Stomata per cm2
(total of upper and 4 16.5 (5)z 20.1 (5)NS 5.0(72) 8.7(72)*
lower surfaces) 12026 15149 6 10.2(72) 16.0(72)** 4.6(5) 6.7(5)*
7 7.5(4) 20.3(4)ns 4.5(4) 7.7(4)ns
Size of leaves in cm2 7 7**
(length x width) 67.64 53.20 Avg 11.4 11.8** 4.7

Total stomata per leaf 8.13 x 105 8.06 x 105 zThe number of single plant readings taken for each average is shown by
the parenthesis.
* and ** significantly greater at the 5% and 1% level respectively from
The upper leaf surface averaged a little more than 10% more values at 85% RH.
stomata than the lower leaf surfaces under both humidity levels.
A calculation of total stomata per leaf, however, demonstra­ individual stomata than on plants grown under 85% RH. Direct
ted that there was a similar number of stomata per leaf on observation of stomatal openings utilizing silicone rubber
plants grown at 85% and at 50% RH. Leaf 4, which had reached impressions was attempted but the impressions were not reliable
full enlargement, had a total of 8.1 X 10$ stomata per leaf indicators.
under both humidity levels (Table 5). Leaves 6 and 7 also had a The diurnal change in stomatal resistance was obtained 3
similar number of stomata per leaf from plants grown under times between 21 and 26 days after seeding for plants grown in
either humidity level even though the leaves were still enlarging liquid culture. Under 85% humidity the resistance of the upper
slowly. The total number of stomata on leaves 6 and 7 was surface remained at about 5 s cm_l for more than half of the

72 J. Amer. Soc. Hort. Sci. 101(1):70—73. 1976.

 RH level in combination with 30% less leaf area should lead to a
large reduction in dry wt accumulation. However, the observed
reduction in dry wt was only 11%, suggesting that photosyn­
thetic rates per unit leaf area of plants growing under the lower
humidity level were not depressed by the higher stomatal
resistance. In fact, photosynthetic rates must have been greater
per unit leaf area on plants grown under low humidity than on
plants grown under high humidity to offset some of the large
differences in leaf area between the plants.
The principal advantage of growing lettuce under high
relative humidities would be the production of marketable
heads in a slightly reduced period of time. These heads would be
larger and have a higher water content than those grown under
HOURS AFTER LIGHTS ON
lower humidities.

Literature Cited
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J. Amer. Soc. Hort. Sci. 101 (1 ):70—73. 1976. 73