Scientia Horticulturae 73 1998.

125136

Pot soil air composition in conditions of high soil moisture and its influence on chrysanthemum growth
Z. Strojny
a b

a,)

, P.V. Nelson b, D.H. Willits

Research Institute of Pomology and Floriculture, 96-100 Skierniewice, Poland Department of Horticultural Science, North Carolina State Uniersity, Raleigh, NC 27695-7625, USA c Department of Biological and Agriculture Engineering, North Carolina State Uniersity, Raleigh, NC 27695-7625, USA Accepted 23 June 1997

Abstract Chrysanthemums were grown in 15.2 cm standard pots in a heavy mix of clay loam soil q sphagnum peat moss 2:1.. A fine texture mix was used to accentuate undesirable gas profiles in the soil. Soil air was analyzed at five depths in the soil profile. In one set of tests, water was applied to the top of the pot at a matrix potential in the center of the soil profile of y5 kPa. The average gas concentrations in soil air in the top and bottom fifths of soil were for O 2 20.0 and 14.5%, for CO 2 0.8 and 2.4%, and for C 2 H 4 0 and 0.08 m l dmy3. Smooth concentration gradients of each gas occurred from top to bottom of the soil profile. The composition of soil air changed greatly during the drying cycle. At soil moisture tensions of y0.7, y2.5, and y5 kPa in the center of the soil profile, the gas concentrations in the lowest fifth of soil were for O 2 9.6, 15.3, and 20.3%, and for CO 2 4.5, 3.5, and 0.6%, respectively. Thus, soil atmospheric conditions for plant growth were poorest immediately after watering and continuously improved up to the time of watering. When pots of chrysanthemum were watered by capillary action from mats, the average concentration of gases in soil air in the lowest fifth of soil were 5.8% O 2 , 3.6% CO 2 , and 0.38 m l dmy3 C 2 H 4 . This gas profile was less desirable for growth than the profile found in top-watered pots. Unlike the situation in top-watered pots, the gas concentrations in mat-watered pots were stable. Roots in top-watered pots were restricted to the upper two thirds of the soil ball, and were distributed evenly in the inner part of the soil and at the periphery. Roots of mat-watered plants developed further down the vertical periphery of the pot than roots of top-watered plants, but they did not grow inside the ball. Chrysanthemum plants were grown through a hole in the side wall of each of five 3.9 cm tall by 15.2 cm diameter plastic rings

Corresponding author. Fax: q48 46 333 088.

0304-4238r98r$19.00 q 1998 Elsevier Science B.V. All rights reserved. PII S 0 3 0 4 - 4 2 3 8 9 7 . 0 0 1 5 6 - 8

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stacked vertically and separated by stainless steel screens that allowed for passage of water but not roots. Water was applied to the top of these cylinder stacks. The largest plants developed in the top ring with progressively smaller plants at lower depths. Plants in the lower two rings developed interveinal chlorosis and did not reach commercial size. q 1998 Elsevier Science B.V.
Keywords: Carbon dioxide; Ethylene; Oxygen; Soil atmosphere; Chrysanthemum; Container media; Watering

1. Introduction A basic condition for root respiration is an adequate supply of O 2 . Oxygen deficit is manifested by wilting, leaf epinasty, chlorosis, poor growth and finally root death Levitt, 1972; Drew and Sisworo, 1977; Kawase, 1981.. Drew 1979., Bradford and Yang 1981. and Glinski and Stepniewski 1985. extensively reviewed studies of plant response to anaerobic conditions. They found that plant reaction to poor soil aeration might be the direct effect of O 2 deficiency, or of an accumulation of respiration products in anoxic or hypoxic conditions in soil or in roots. According to these studies, some reduced ions NO 2y, Mn2q, S 2y, Fe 2q . and intermediate products in microbial carbon metabolism, such as volatile fatty acids, phenolic acids and various hydrocarbons including ethylene can accumulate in the soil to harmful levels for plants. It is a matter of discussion how extensive O 2 deficiency needs to be before negative plant effects occur. Investigations of these problems have been carried out for many years. They were conducted mainly under field conditions. Not much is known about substrates and container conditions in the greenhouse. According to numerous works reviewed by Heiskanen 1993., adequate substrate aeration is of critical importance for plants grown in containers. It can be achieved in container substrate that is saturated to container capacity when air space is at least 10% of total substrate volume Bugbee and Frink, 1986.. Air exchange in soil can be limited by fine texture, compactness, and high water content leading to lower O 2 and higher CO 2 concentrations. It is difficult to define the critical concentrations of O 2 and CO 2 in soils for plants because plant requirements can change in accordance with the rate of root respiration, which reflects the root activity. According to Kramer 1969., the usual concentration of O 2 in soil air can be low enough to harm plants, but, the CO 2 level is generally not high enough to be injurious. The objectives of this study were to determine concentrations of O 2 , CO 2 , and to a lesser extent C 2 H 4 throughout the root ball profile in pots of a heavy soil mix during chrysanthemum production, effects of water application to the top of the pot vs. through capillary mats on soil air composition, and the growth response of chrysanthemum to soil conditions at five depths in top-watered pots.

2. Material and methods Three experiments were conducted in a glass greenhouse. In the first, chrysanthemums Cymbals were grown from March 27 to a market stage on June 9, in 15.2 cm

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Fig. 1. Moisture release curve for the 2:1 clay loam soilqsphagnum peat moss soil mix used in the three experiments of this study. Soil mix physical properties included a bulk density of 0.91, total porosity of 56%, and container water capacity of 51.7%.

diameter standard depth plastic pots 14.4 cm deep. in a mix of clay loam soil q sphagnum peat moss 2:1. Fig. 1.. There were 5 plants per pot. Incandescent light was applied from 2200 to 0200 h for the first 12 days followed by the application of black shade cloth from 1800 to 0800 h until color was well developed in buds. The terminal 2 cm of each plant was removed at 12 days. All nutrients were applied in a pre-plant manner. Peat moss was amended with 8 g dolomitic limestonerl prior to mixing with soil. The soil mix was amended with 1 g single superphosphate plus 6 g of Osmocote 14.0-6.1-11.6 slow-release fertilizer mini-capsule [Link] Scott-Sierra, Marysville, OH.. Soil air was drawn weekly, regardless of soil moisture tension, from sample cells at five depths, each located 2 cm inside the pot wall. All samples on a given date were drawn at the same time, but the time was not set with respect to a given point in the drying cycle. Sample cells were positioned spirally around the pot in order to increase the distance between each, and in the middle depth at additional distances of 0.5 and 6.5 cm from the pot wall Fig. 2a.. Water was applied to the top of the pots to the point of full saturation when the matrix potential reached y5 kPa, as measured at the center of the soil profile by tensiometers.

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Fig. 2. Distribution of air sampling cells in soil ball numbers show the distance from pot bottomrwall in cm. for a. Experiments 1 and 2 and b. Experiment 3.

Sample cells Fig. 3. had a volume of 1.5 cm3 and consisted of a plastic mesh cylinder, a womans hair curler, covered with one layer of cheese cloth, that measured 1 cm in diameter by 2 cm in length. A glass tube of 2 mm inside diameter extended from the cavity of the sample cell to a rubber septum imbedded in a hole in the pot wall. Gas was sampled with a 1 cm3 syringe inserted through the septum. Oxygen and CO 2 were analyzed weekly while C 2 H 4 was analyzed twice at four and six weeks. There were six single pot replications. Soil air was analyzed in a gas chromatograph Fisher Gas Partitioner Model 1200.. The results were tested by an analysis of variance for two-factor design.

Fig. 3. Diagram of air sampling cell: 1. plastic mesh cylinder with openings circa 3=1.5 mm, 2. glass tube with inner diameter of 2 mm, 3. rubber stopper, 4. rubber septum, 5. pot wall.

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In the second experiment, similar pots were used with five chrysanthemum Yellow Mandalay plants. One set of six pots was sub-irrigated from a capillary mat kept constantly moist with tap water, while a second set of six pots was watered at the top of the pot at a soil water potential of y5 kPa in the center of the soil profile. Plants were potted on June 24 and the terminal 2 cm of each was removed 14 days later. Natural long day conditions prevailed for 14 days followed by black cloth application daily from 1800 to 0700 h. The experiment was terminated on Aug. 1, when plants were five weeks old. The same pre-plant fertilization procedures were used in this experiment as in the first experiment. Soil air samples were taken from the three lowest locations described in Experiment 1. Soil air samples were drawn weekly from the second through fifth consecutive weeks in the case of irrigation by capillary mat, and during the third, fifth and sixth weeks in the case of water applied to the top. In the latter case, O 2 and CO 2 concentrations were determined within each of these three water cycles 2 h after watering when excess gravitational water had drained y0.7 kPa, in the middle of the watering cycle at y2.5 kPa, and shortly before watering at y5 kPa. Ethylene concentration was measured twice, at four and six weeks. Data from mat-watered plants were statistically tested by a one-way analysis of variance based on four replications in time sampling dates. for O 2 and CO 2 , and two replications in time for C 2 H 4 . Data from top-watered plants were tested by an analysis of variance for two factors based on three replications in time three watering cycles.. The third experiment was conducted twice to investigate the extent of plant development possible when plant roots were restricted to each of five vertically stacked soil zones. A PVC plastic pipe 15.2 cm in diameter and 14.5 cm in length was cut into five equal rings. These were stacked vertically with a stainless steel screen, with openings of 0.16 mm, separating each. The screens and pipe rings were connected together with latex cement. A gas sampling tube was installed in each ring as shown in Fig. 2b. Water could pass through the screens, while roots could not. A single-rooted chrysanthemum Cymbals cutting was planted on 17 June in Experiment 3a, and on 4 September in Experiment 3b through a hole in the side wall of each ring by the following procedure. A hole was drilled through the center of a rubber stopper. The stopper was cut into two halves from top to bottom. The two stopper pieces were placed around the stem of a plant, and lanoline paste was used to seal any remaining space between the stem and the rubber. Finally, the stopper was inserted snugly into a hole in the pot wall. The holes were arranged in a semi-spiral covering an arc of 1208, so one plant was not situated over another, and all had a south exposure in the greenhouse Fig. 4.. The terminal part above sixth leaf of each plant was removed 14 days after planting. Plants were grown under natural long days for 11 days, and were shaded daily from 1800 to 0700 h thereafter. Water or fertilizer was applied to the top cylinder when the water potential in the center of the middle ring reached y5 kPa. Six hundred milliliters of water or fertilizer was applied to each cylinder to achieve a leaching percentage of 20 to 30%. Once a week, fertilizer consisting of 600 mg N q 130 mg phosphate-Pq 506 mg K derived from NH 4 NO 3 , KNO 3 , and NH 4 . 2 HPO4 was applied. Prior to planting, the soil mix was amended with 1.5 g single superphosphate plus 6 g dolomitic limestone, plus 3 g Esmigran micronutrient mix Scotts-Sierra Horticultural Products, Marysville,

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Fig. 4. Six-week-old chrysanthemum plants, each grown through a hole in the side wall of one of five vertically stacked cylinders in Experiment 3. Cylinders were separated by stainless-steel screens that permitted the passage of water but not roots. a. Plants growing in the cylinder stack. b. Plants removed from the cylinders. In both photos, plants from left to right were grown in progressively higher cylinders from bottom to top.

OH. per liter of soil. There were six single column replications, and the experiment was conducted twice. At the termination of the experiments on 31 July and 10 October, plants were cut at the stopper. The height of each plant, the number of leaves on each plant, and the fresh weight of each plant was measured. Results were tested by analysis of variance design for two factors. Means from all experiments were evaluated using Duncans multiple-range test for 5% level of significance.

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3. Results and discussion 3.1. Experiment 1 The O 2 concentration was lowest at the bottom of the soil profile Table 1.. The average concentration at this depth was 14.5%, but fluctuations between measurements over weeks were high. The fluctuations were suspected to be due to differences in soil moisture tension from week to week when samples were taken. The lowest concentration occurred during weeks 2 through 5 when the weather was cloudy, and young plants did not absorb much water, both causing the soil to remain moist. The O 2 concentration increased, and its fluctuation decreased as the distance to the soil surface decreased. Near the soil surface, the O 2 concentration was only a little lower than in ambient air. There were no differences in O 2 concentration depending on the distance from the side of soil ball in the middle layer. The opposite trend was found in CO 2 concentration Table 2.. The concentrations were highest in the two positions closest to the bottom y2.4% on average. The concentrations decreased with increasing proximity to the upper soil surface. But even in the top layer, it was much higher 0.8% on average. than in ambient air 0.035%.. Fluctuation over weeks was also high, but not as high as for O 2 . Concentration did not change with distance from the pot wall in the middle layer. Ethylene was not found in the two top layers but was found in the lower layers Table 3.. Ethylene concentration increased with greater depth in the soil to a maximum average concentration of 0.133 m l dmy3 in the bottom layer during the fourth week. According to Abbeles 1982., a concentration above 0.1 m l dmy3 can be harmful for plants. The results of this experiment indicated that the soil atmosphere in the lower half

Table 1 Mean O 2 concentration %. at the end of each of 10 weeks, mean O 2 concentration of all weeks, and the range of O 2 concentrations for all weeks and replications in soil air in 15.2 cm top-watered pots of chrysanthemums in Experiment 1 Week Distance from pot bottomrwall cm. 1.5r2 1 2 3 4 5 6 7 8 9 10 Mean Range 20.3 8.3 10.5 5.7 14.2 18.5 17.7 15.7 16.3 18.0 14.5 3.420.9 4.3r2 20.6 15.7 16.9 16.8 19.2 18.9 18.1 17.9 18.4 19.1 18.2 5.921.0 7.1r0.5 20.9 18.7 18.6 17.7 19.6 19.2 18.7 17.9 18.6 19.1 18.9 15.321.0 7.1r2 20.8 18.2 18.3 16.6 19.1 18.8 18.1 18.1 18.3 19.5 18.6 13.221.0 7.1r6.5 21.0 18.2 19.7 16.8 17.9 18.7 18.7 18.0 17.9 19.4 18.6 13.721.0 9.9r2 21.0 19.2 19.5 16.9 19.7 19.6 18.6 18.5 18.9 19.4 19.2 13.821.0 12.7r2 21.0 19.8 19.9 17.3 19.5 20.1 19.7 20.4 20.6 20.8 20.0 14.521.0 LSD 0.05

2.04

0.65

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Table 2 Mean CO 2 concentration %. at the end of each of 10 weeks, mean CO 2 concentration of all weeks, and the range of CO 2 concentrations for all weeks and replications in soil air in a 15.2 cm top-watered pots of chrysanthemums in Experiment 1 Week Distance from pot bottomrwall cm. 1.5r2 1 2 3 4 5 6 7 8 9 10 Mean Range 1.21 3.24 3.71 1.58 1.97 2.67 2.66 2.86 2.21 1.61 2.37 0.485.95 4.3r2 0.86 4.60 2.50 2.67 2.03 2.99 2.95 2.57 1.97 1.66 2.48 0.525.47 7.1r0.5 0.82 3.22 1.57 1.59 1.74 2.70 2.66 2.38 1.90 1.48 2.01 0.464.54 7.1r2 0.82 3.12 1.52 1.68 1.56 2.63 2.62 2.30 1.79 1.52 1.96 0.474.36 7.1r6.5 0.87 3.30 1.66 1.92 1.69 2.83 2.66 2.59 1.89 1.56 2.10 0.444.55 9.9r2 0.78 2.74 1.30 1.38 1.11 1.97 2.12 2.06 1.35 1.23 1.61 0.434.56 12.7r2 0.62 1.70 0.91 0.66 0.40 1.17 1.01 0.64 0.48 0.46 0.80 0.144.34 LSD 0.05

0.784

0.248

of the root ball could be unfavorable for plant growth. This was further suggested by the results of other studies Stolzy et al., 1961; Williamson, 1968; Huck, 1970.. To test this suspicion, the following experiments were conducted. 3.2. Experiment 2 The mean gas composition of air over weeks 2 through 5 in the lowest layer of soil in pots of chrysanthemum watered on mats was low in O 2 , 5.75%, high in CO 2 , 3.63%, and high in C 2 H 4 , 0.379 m l dmy3 Table 4.. In the two soil levels above the bottom level, the O 2 concentration was considerably higher and close to the ambient air concentration. CO 2 and C 2 H 4 concentrations were progressively lower in the second and third layers above the pot bottom. Most notable was the lack of a significant change in gas content of the soil air during the four weeks of sampling. In the top-watered soil treatment, there were large changes in soil air composition during the watering cycle Table 5.. In samples taken shortly before water application at

Table 3 Mean C 2 H 4 concentration m l dmy3 . at the end of weeks 4 and 6 and the range of C 2 H 4 concentrations for both weeks and all replications in soil air in 15.2 cm top-watered pots of chrysanthemums in Experiment 1 Week Distance from pot bottomrwall cm. 1.5r2 4 6 Range 0.133 0.022 00.397 4.3r2 0.056 0.009 00.118 7.1r0.5 0.042 0.000 00.143 7.1r2 0.018 0.007 00.041 7.1r6.5 0.024 0.000 00.042 9.9r2 0.000 0.000 12.7r2 0.000 0.000 0.0446 LSD 0.05

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Table 4 Mean O 2 , CO 2 , and C 2 H 4 concentrations over weeks 2 through 5 and the range of concentrations of these gases for all weeks and replications in soil air in 15.2 cm pots of chrysanthemums watered on a capillary mat in experiment 2 Distance from pot bottom cm. 1.5 means Range 4.3 means Range 7.1 means Range LSD 0.05 for means O 2 %. 5.75 3.329.27 18.10 14.519.8 19.08 16.520.6 1.523 CO 2 %. 3.63 1.775.01 2.19 1.073.88 1.56 0.942.48 0.681 C 2 H 4 m l dmy3 . 0.379 0.0020.666 0.033 0.0000.087 0.020 0.0000.045 0.1607

y5 kPa, the O 2 concentration at all points of sampling was similar to ambient air. Concentration of CO 2 was much higher 0.60 to 0.67%. than in ambient air, but low compared to later concentrations in soil. The situation dramatically changed 2 h after watering. The O 2 concentration in the two lowest fifths of the soil profile dropped to 10% or lower and in the middle fifth to about 15%. The CO 2 concentrations increased to over 4% in the lower three fifths of soil with some individual replications exceeding 7%. When the soil dried to y2.5 kPa, the gas concentrations changed to levels intermediate between just after watering and y5 kPa. Ethylene was found only in samples taken

Table 5 Mean O 2 and CO 2 concentrations %. for weeks 2, 4, and 5 combined and the range of concentrations of these gases for all three weeks and replications in soil air in 15.2 cm top-watered pots of chrysanthemums just before watering y5 kPa., 2 h after watering y0.7 kPa., and at the middle of the water cycle y2.5 kPa. in Experiment 2 a Distance from pot bottom cm. O2 1.5 means Range 4.3 means Range 7.1 means Range CO2 1.5 means Range 4.3 means Range 7.1 means Range
a

Before watering 20.3c 19.521.0 20.5c 20.021.0 20.4c 18.520.9

After watering 9.6a 4.417.3 10.0a 3.516.0 15.6b 9.217.7

Middle of watering cycle 15.3b 9.318.9 17.9bc 14.419.8 18.53bc 17.119.7

0.60a 0.221.34 0.67a 0.341.38 0.63a 0.321.21

4.47de 2.296.28 5.21e 3.137.31 4.58de 3.237.01

3.53cd 1.655.42 2.27bc 1.544.56 2.38b 1.393.94

Means with the same letters do not differ at the level of significance 0.05 according to Duncans multiple-range test.

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shortly after watering. Concentrations were highest in the lowest part of the soil profile, and only trace amounts appeared in the higher positions. These changes in top-watered pots indicate that measured gas values reflect the composition of air contained in macropores. Diffusion of air into the soil brings the soil air composition to a state not much different than that in ambient air. However, this macropore gas probably differs greatly from the composition of air trapped in micropores. Infiltrating water during its application apparently displaced air contained in micropores into the macropores, substantially changing the composition of air in these surrounding macropores. Overall, mat-watered soil at all times and top-watered soil just after watering contained very deficient amounts of O 2 and possibly excessive amounts of CO 2 and C 2 H 4 , especially in the lower part of the pot. In both watering schemes at these stated times, the soil atmosphere in the lowest three soil zones appeared to be undesirable for growth. Comparatively speaking, the gas composition in the second and third layer from the pot bottom was always better in the mat-watered pots than in the top-watered pots shortly after watering. This assessment is supported by the observation that plant roots in pots irrigated by capillary mat were evenly distributed from top to bottom at the side surface of the root ball, but only extended about halfway down the pot wall in top-watered pots Fig. 5.. It was also observed that roots developed laterally across the soil profile in top-watered plants but remained closer to the pot periphery in mat-watered pots. Heiskanen 1993. observed that under wet conditions, roots tended to spread to the periphery of the container. Also, Biran and Eliassaf 1980. noticed that in the poorly aerated central part of the root ball, there was a considerable decrease in root quantity. Perhaps the driest period prior to watering in top-watered pots fostered root development in the center of the root ball.

Fig. 5. Distribution of roots in the soil ball after 38 days of chrysanthemum growth left. from a pot watered by a capillary mat and right. a pot watered from the top at a soil moisture tension of y5 kPa in the center of the ball in Experiment 2.

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Table 6 Mean stem length, number of leaves per plant, and plant fresh weight of chrysanthemum plants restricted to each of five vertically stacked zones in a top-watered 14.5 cm tall by 15.2 cm diameter column see Fig. 2b. a Zone cm from pot bottom. 0.02.9 2.95.8 5.88.7 8.711.6 11.614.5
a

Stem length cm. 7.0a 15.7b 17.9c 19.0c 19.8c

No. of leaves per plant 48a 79b 86b 102c 113d

Fresh weight g. 10.4a 28.8b 41.2c 54.3d 68.6e

Symptomsb qq q y y y

Means with the same letters do not differ at the level of significance 0.05 according to Duncans multiple-range test. b qq: well-developed interveinal leaf chlorosis; q: light leaf chlorosis; y: no symptoms. All values are the combined mean of measurements from Experiments 3a and 3b.

3.3. Experiment 3 The composition of soil air in each of the five rings data not shown. was similar to that in top-watered pots in Experiment 2 Table 5.. Differences in the growth of plants, each with roots restricted to one depth of soil, occurred as a consequence of soil depth early in the crop and became more pronounced with time Table 6.. Plant size decreased with closer proximity to the base of the pot. At the end of the crop, plants in the lower two soil zones did not reach marketable quality, while plants in the middle zone were of questionable marketable quality. The number of leaves per plant and plant weight increased continually from the bottom to the top soil zones. Stem length was shorter in the lower two soil zones than in the three top zones. Plants in the lowest soil zone developed extensive interveinal chlorosis, while those in the next zone up developed a moderate level of interveinal chlorosis. Plants in the three top zones had a normal green color. A visual assessment of roots indicated that the size of the root system was in proportion to the size of the shoot.

4. Conclusions The suitability of gas composition for plant growth decreased from the top to the bottom of the soil profile in 15.2 cm diameter pots containing plants. The most adverse gas contents found in the lowest 20 % of the soil were 3% O 2 , 6% CO 2 , and 0.67 m l dmy3 C 2 H 4 . Levels of C 2 H 4 above the 0.1 m l dmy3 upper critical level for plants occurred in each of the lower three fifths of the soil profile. When plants were restricted to a given 20% of the vertical soil profile, growth was unacceptable in the lower three fifths of the profile. Likewise, when plants were planted in the top of the pot, roots did not grow well in the lower portion of the soil column. These growth results confirmed the former assessment of soil gas content. The gas content of the lower three fifths of soil was poor and remained reasonably constant over time when plants were watered by capillary mats. When watered from the top, soil gas composition was more adverse for

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plant growth shortly after watering than it was in mat-watered soil profiles. However, gas composition improved in top-watered soil during drying to a point just before watering, where it was much superior to that in mat-watered soil. State-of-the-art substrates used today could be expected to have gaseous concentration gradients, although not as extreme as those encountered in the heavy soil used in this study. Undoubtedly, if current substrates were watered too frequently, suppression of root growth would be anticipated in the lower horizons of the soil profile.

Acknowledgements The authors wish to thank Dr. Sylvia Blankenship for advice and help in analysis of gas samples.

References
Abbeles, F.B., 1982. Ethylene as an air pollutant. Agric. Fores. Bull. 5 1., 412. Biran, I., Eliassaf, A., 1980. The effect of container size and aeration conditions on growth of roots and canopy of woody plants. Sci. Hort. 12, 385394. Bradford, K.J., Yang, S.F., 1981. Physiological responses of plants to waterlogging. Hort. Sci. 16, 2530. Bugbee, G.J., Frink, C.R., 1986. Aeration of potting media and plant growth. Soil Sci. 141, 438441. Drew, M.C., Sisworo, E.J., 1977. Early effects of flooding on nitrogen deficiency and leaf chlorosis in barley. New Phytol. 79, 567571. Drew, M.C., 1979. Plant responses to anaerobic conditions in soil and solution culture. Curr. Adv. Pl. Sci. 36, 114. J. Glinski, W. Stepniewski, 1985. Soil Aeration and its Role for Plants. CRC Press, Boca Raton, FL. Heiskanen, J., 1993. Favorable water and aeration conditions for growth media used in containerized tree seedling production: a review. Scand. J. For. Res. 8, 337358. Huck, M.G., 1970. Variation in taproot elongation rate as influenced by composition of the soil air. Agron. J. 62, 815818. Kawase, M., 1981. Anatomical and morphological adaptation of plants to waterlogging. Hort. Sci. 16, 3034. P.J. Kramer, Plant and Soil Water Relationships. McGraw-Hill, New York, 1969. J. Levitt, Responses of Plants to Environmental Stress. Academic Press, New York, 1972. Stolzy, L.H., Letey, J., Szuszkiewicz, T.E., Lunt, O.R., 1961. Root growth and diffusion rates as functions of oxygen concentration. Soil Sci. Soc. Am. Proc. 25, 465467. Williamson, R.E., 1968. Influence of gas mixtures on cell division and root elongation of broad bean Vicia faba L. Agron. J. 60, 317321.