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G Aerial dispersal and drying of Peronospora tabacina conidia in ...

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700 Applied Physical Sciences: Aylor <strong>and</strong> Taylor<br />

Table 1. Average time for leaves to become half dry (t12) <strong>and</strong><br />

fully dry (tf)<br />

Date Wall Location<br />

t1/2P<br />

m<strong>in</strong><br />

tf,<br />

m<strong>in</strong><br />

May 22 Down<br />

Up<br />

N1<br />

Si<br />

163<br />

160<br />

178<br />

179<br />

Down N3 115 169<br />

Up S3 123 174<br />

Down N5 135 170<br />

Up S5 138 168<br />

July 1 Down N1 191 248<br />

Up Si 164 230*<br />

Down N2 171 236<br />

Up S2 170 230<br />

Down N4 181 246<br />

Up S4 160 225*<br />

Down N5 175 243<br />

Up S5 160 235<br />

Measurements were made at six locations after 0700 EST May 22<br />

<strong>and</strong> at eight locations after 0800 EST July 1. Locations N1-N5 <strong>and</strong><br />

S1-S5 are shown <strong>in</strong> Fig. 1.<br />

* Values for which tf was significantly less with the wall raised.<br />

4 to about 12 m, but at 20 m Rc was still about 1.8. The percentage<br />

<strong>of</strong> spores escap<strong>in</strong>g the tent <strong>in</strong>creased from 0.005% to<br />

about 0.46%. When Kz <strong>in</strong>creased exponentially, RC did not become<br />

greater than 1.0 until 18 m from the source, <strong>and</strong> at 20 m<br />

it was only 1.15. Meanwhile, the percentage <strong>of</strong> spores potentially<br />

escap<strong>in</strong>g the tent <strong>in</strong>creased to fully 22%.<br />

Dry<strong>in</strong>g. The times t1/2 <strong>and</strong> tf for leaves to become half <strong>and</strong><br />

fully dry are shown <strong>in</strong> Table 1. Generally, t1/2 was 70-90% <strong>of</strong><br />

On May 22, w<strong>in</strong>d just outside the tent was north-northwest<br />

<strong>and</strong> 0.3-1.2 m/sec. From 0735 to 1000 EST, air temperature<br />

<strong>in</strong>creased from 10'C to 24TC, while relative humidity decreased<br />

from 87% to 51%. The tf was about 170 m<strong>in</strong>. There were no<br />

significant differences <strong>in</strong> the time for leaves to dry whether walls<br />

were up or down.<br />

On July 1, w<strong>in</strong>d was north-northwest <strong>and</strong> calm to 1.4 m/sec.<br />

From 0818 to 1155 EST, air temperature <strong>in</strong>creased from 20TC<br />

to 270C while relative humidity decreased from 86% to 72%.<br />

The tf was about 230 m<strong>in</strong>. There were slight differences <strong>in</strong> the<br />

time for leaves to dry, depend<strong>in</strong>g on whether the wall was up<br />

or down. In two <strong>of</strong>four locations <strong>in</strong>side the tent, the leaves dried<br />

a significant 7-8% faster with the walls up.<br />

To generalize, we calculated the effect <strong>of</strong> rais<strong>in</strong>g the walls on<br />

the time for leaves to dry. Although the <strong>in</strong>itial water on the leaf,<br />

the radiation absorbed by the leaf, <strong>and</strong> leaf size were not known<br />

precisely, these factors were not affected by wall position <strong>and</strong><br />

calculation provided a valid comparison <strong>of</strong> walls up <strong>and</strong> down.<br />

Rais<strong>in</strong>g the end walls can <strong>in</strong>crease air temperature, decrease<br />

relative humidity <strong>and</strong> <strong>in</strong>crease w<strong>in</strong>d speed <strong>in</strong>side the tent. Depend<strong>in</strong>g<br />

on the size <strong>of</strong> the plants, w<strong>in</strong>d speed just <strong>in</strong>side the tent<br />

can be <strong>in</strong>creased 40-70% by rais<strong>in</strong>g the walls. The air is generally<br />

about 1C warmer <strong>and</strong> about 5-7% drier outside than <strong>in</strong>.<br />

Thus, rais<strong>in</strong>g the walls could at most change w<strong>in</strong>d, temperature,<br />

Proc. Nad Acad. Sci. USA 79 (1982)<br />

<strong>and</strong> humidity <strong>in</strong>side the tent by these amounts, <strong>and</strong>, us<strong>in</strong>g these<br />

values, we calculated an upper limit on the effect <strong>of</strong> rais<strong>in</strong>g the<br />

walls. Penman's method for estimat<strong>in</strong>g evaporation from an extensive<br />

wetted surface (1) <strong>in</strong>dicates that these changes would<br />

make essentially no difference <strong>in</strong> the time for the entire crop<br />

to dry. We calculated the <strong>dry<strong>in</strong>g</strong> <strong>of</strong> an <strong>in</strong>dividual upper leaf <strong>and</strong><br />

found that the maximal changes <strong>in</strong> w<strong>in</strong>d, temperature, <strong>and</strong> humidity,<br />

had they occurred, might have caused upper leaves to<br />

dry about 30 m<strong>in</strong> faster on May 22 <strong>and</strong> about 40 m<strong>in</strong> faster on<br />

July 1.<br />

Germ<strong>in</strong>ation. Would a 30-m<strong>in</strong> shorter <strong>dry<strong>in</strong>g</strong> time affect the<br />

fungus? The critical stage <strong>in</strong> the disease cycle is germ<strong>in</strong>ation <strong>of</strong><br />

spores <strong>and</strong> the penetration <strong>of</strong> the leaf. These stages require<br />

water, whereas the subsequent stages can proceed <strong>in</strong> dry leaves.<br />

Thus the significance <strong>of</strong> 30 m<strong>in</strong> can be judged by the slope <strong>of</strong><br />

the curve relat<strong>in</strong>g percent germ<strong>in</strong>ation to time after wett<strong>in</strong>g<br />

spores. Between 150C <strong>and</strong> 30TC, germ<strong>in</strong>ation is most rapid dur<strong>in</strong>g<br />

the first 120 m<strong>in</strong> after wett<strong>in</strong>g <strong>and</strong> considerably slower<br />

thereafter (16). Between 0 <strong>and</strong> 120 m<strong>in</strong> a difference <strong>of</strong> 30 m<strong>in</strong><br />

could <strong>in</strong>crease germ<strong>in</strong>ation by 16-18%. Thus, where leaves dry<br />

dur<strong>in</strong>g the first 120 m<strong>in</strong> after the spores are first wet, a 30 m<strong>in</strong><br />

faster <strong>dry<strong>in</strong>g</strong> might significantly reduce germ<strong>in</strong>ation. Usually,<br />

however, the spores <strong>and</strong> leaves will be wet considerably longer<br />

than 120 m<strong>in</strong>. Between 120 <strong>and</strong> 240 m<strong>in</strong> a difference <strong>of</strong> 30 m<strong>in</strong><br />

could <strong>in</strong>crease germ<strong>in</strong>ation by 2-6%. By 240 m<strong>in</strong> germ<strong>in</strong>ation<br />

has reached its maximum <strong>and</strong> more time would not matter.<br />

Thus, 30 m<strong>in</strong> faster <strong>dry<strong>in</strong>g</strong> would generally decrease <strong>in</strong>fection<br />

less than 6%, a small matter.<br />

Rais<strong>in</strong>g the end walls affects the w<strong>in</strong>d <strong>and</strong> thus the <strong>dispersal</strong><br />

<strong>of</strong> spores from lesions near the edge <strong>of</strong> the crop but will have<br />

little effect on <strong>dry<strong>in</strong>g</strong>. Blue mold <strong>of</strong>ten starts near the edge <strong>of</strong><br />

a tent where hedges or sheds shade the plants from the morn<strong>in</strong>g<br />

sun <strong>and</strong> where wetter, cooler conditions encourage the fungus.<br />

Thus, rais<strong>in</strong>g the walls could <strong>in</strong>crease the <strong>in</strong>oculum deposited<br />

on plants well <strong>in</strong>to the tent <strong>and</strong> <strong>in</strong>crease the number <strong>of</strong> spores<br />

escap<strong>in</strong>g the tent <strong>and</strong> spread<strong>in</strong>g the disease afar.<br />

The authors thank P. A. Halbert <strong>and</strong> K. T. Paw U for technical assistance<br />

<strong>and</strong> G. J. S. Ross <strong>of</strong> Rothamsted Experimental Station for help<br />

with the statistical analysis <strong>of</strong> leaf <strong>dry<strong>in</strong>g</strong>.<br />

1. Waggoner, P. E., Pack, A. B. & Reifsnyder, W. E. (1959) Bullet<strong>in</strong><br />

626 (Connecticut Agric. Exper. Station, New Haven, CT).<br />

2. Lucas, G. B. (1980) Science 210, 147-153.<br />

3. Smith, F. B. & Hay, J. S. (1961) Quart.J. R. Met. Soc. 87, 82-101.<br />

4. Sutton, 0. G. (1953) Micrometeorology (McGraw-Hill, New<br />

York).<br />

5. Thom, A. S. (1975) <strong>in</strong> Vegetation <strong>and</strong> the Atmosphere (Academic,<br />

London), Vol 1.<br />

6. Belot, Y., Baille, A. & Delmas, J.-L. (1976) Atmos. Environ. 10,<br />

89-98.<br />

7. Roache, P. J. (1972) Computational Fluid Dynamics (Hermosa,<br />

Albuquerque, NM).<br />

8. Legg, B. J. & Powell, F. A. (1979) Agric. Meteorol. 20, 47-67.<br />

9. Gregory, P. H. (1973) The Microbiology <strong>of</strong> the Atmosphere<br />

(Wiley, New York).<br />

10. May, K. R. & Clifford, R. (1967) Ann. Occup. Hyg. 10, 83-95.<br />

11. Ross, G. J. S. (1980) Maximum Likelihood Program (Rothamsted<br />

Experimental Station, Harpenden, Engl<strong>and</strong>).<br />

12. Raschke, K. (1956) Planta 48, 200-238.<br />

13. Monteith, J. L. (1973) Pr<strong>in</strong>ciples <strong>of</strong> Environmental Physics<br />

(American Elsevier, New York).<br />

14. Waggoner, P. E. (1975) <strong>in</strong> Vegetation <strong>and</strong> the Atmosphere (Academic,<br />

London), Vol. 1.<br />

15. Turner, N. C. & Begg, J. E. (1973) Plant Physiol. 51, 31-36.<br />

16. Cruickshank, I. A. M. (1961) Aust. J. Biol. Sci. 14, 58-65.

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