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Factors involved in ice nucleation and propagation ... - Landbunadur.is

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44 BÚVÍSINDI<br />

Figure 2. Top panel: Upper (top left) <strong>and</strong> lower<br />

(top right) surface of a bean (Phaseolus vulgar<strong>is</strong>)<br />

leaf show<strong>in</strong>g the presence of stomates on the abaxial<br />

surface. Middle panel: Infrared images of bean<br />

leaf with drops of water conta<strong>in</strong><strong>in</strong>g <strong>ice</strong>-<strong>nucleation</strong>-active<br />

(INA) bacteria (Pseudomonas syr<strong>in</strong>gae,<br />

stra<strong>in</strong> Cit7). On the left, drops have frozen but<br />

leaves have not, while on the right the leaf with<br />

the frozen droplet on the abaxial surface has frozen<br />

but the leaf with the frozen droplet on the<br />

adaxial surface rema<strong>in</strong>s unfrozen. Bottom panel:<br />

Infrared images of abaxial surface of bean leaves<br />

with drops of water conta<strong>in</strong><strong>in</strong>g INA bacteria on<br />

the surface. The leaf on the left has a layer of<br />

silicone grease under the droplet of INA bacteria.<br />

On the left, droplets have frozen but leaves have<br />

not while on the right, the leaf with the silicone<br />

grease subtend<strong>in</strong>g the droplet has not frozen while<br />

the leaf with only the droplet of INA bacteria has<br />

frozen. Th<strong>is</strong> <strong>in</strong>dicates that <strong>ice</strong> must physically<br />

grow through a stomate or crack <strong>in</strong> the cuticle <strong>in</strong><br />

order to <strong>in</strong>itiate freez<strong>in</strong>g of the leaf. (Repr<strong>in</strong>ted<br />

from W<strong>is</strong>niewski <strong>and</strong> Fuller, 1999, with perm<strong>is</strong>sion).<br />

2. mynd. Efsta röð: Efra borð (til v<strong>in</strong>stri) og neðra<br />

borð (til hægri) laufblaðs baunaplöntu (Phaseolus<br />

vulgar<strong>is</strong>) sýnir loftauga á ásfrálægu yfirborði. Miðröð:<br />

Innrauð mynd af baunablaði með vatnsdropa<br />

sem <strong>in</strong>niheldur ískr<strong>is</strong>tallamynd<strong>and</strong>i (INA) bakteríu<br />

(Pseudomonas syr<strong>in</strong>gae, stofn Cit7). V<strong>in</strong>stra meg<strong>in</strong><br />

hefur drop<strong>in</strong>n frosið, en laufblaðið ekki, og hægra<br />

meg<strong>in</strong> hefur laufblaðið á ásfrálæga yfirborð<strong>in</strong>u<br />

frosið, en laufblaðið á ásaðlæga yfirborð<strong>in</strong>u er<br />

ófrosið. Neðsta röð: Innrauð mynd af ásaðlæga<br />

ter can freeze on the upper surface of these<br />

plants <strong>and</strong> the plant will cont<strong>in</strong>ue to supercool.<br />

When external <strong>ice</strong> does <strong>in</strong>duce the plant<br />

to freeze it <strong>is</strong> by the growth of <strong>ice</strong> through a<br />

stomatal open<strong>in</strong>g on the abaxial surface. In<br />

herbaceous plants, the cuticle <strong>is</strong> not an effective<br />

barrier, or there are sufficient avenues of<br />

<strong>in</strong>gress that allow <strong>ice</strong> to readily propagate from<br />

either the upper or lower surface. Provid<strong>in</strong>g a<br />

barrier of silicone grease sufficiently prevents<br />

external <strong>ice</strong> from <strong>in</strong>duc<strong>in</strong>g herbaceous plants<br />

to freeze. To test the hypothes<strong>is</strong> that a hydrophobic<br />

barrier can prevent plants from freez<strong>in</strong>g,<br />

an emulsion of hydrophobic kaol<strong>in</strong> (Englehard,<br />

Inc.) was applied to the surface of leaves<br />

(Figure 3) or whole 4–6 week old tomato plants<br />

prior to application of an extr<strong>in</strong>sic nucleat<strong>in</strong>g<br />

agent (Cit7 stra<strong>in</strong> of Ina + Pseudomonas syr<strong>in</strong>gae).<br />

Results <strong>in</strong>dicated that dry, young tomato<br />

plants can supercool to as low as –6°C<br />

whereas plants hav<strong>in</strong>g a s<strong>in</strong>gle droplet of Cit7<br />

would freeze at –1.5 to –2.5°C. Application<br />

of the hydrophobic barrier blocked the effect<br />

of Cit7 <strong>and</strong> allowed the plants to supercool to<br />

–6°C, despite the presence of frozen droplets<br />

on leaf surfaces.<br />

THE INITIAL FREEZING PROCESS AND<br />

SUBSEQUENT ICE PROPAGATION<br />

Ice <strong>nucleation</strong>, when it occurs at warm temperatures<br />

<strong>in</strong> herbaceous plants, <strong>is</strong> a two step<br />

process. In the first step, only water that <strong>is</strong><br />

present along the surface of cells <strong>is</strong> <strong>in</strong>duced<br />

to freeze. Th<strong>is</strong> can be seen as a small exothermic<br />

response after which the plant t<strong>is</strong>sue quickly<br />

yfirborð<strong>in</strong>u með dropa með ískr<strong>is</strong>tallamynd<strong>and</strong>i<br />

bakteríu á yfirborð<strong>in</strong>u. Laufblaðið til v<strong>in</strong>stri er<br />

með lag af silikonfeiti undir dropanum. Til v<strong>in</strong>stri<br />

hafa droparnir frosið, en laufblaðið ekki, og hægra<br />

meg<strong>in</strong> hefur drop<strong>in</strong>n á silikon<strong>in</strong>u ekki frosið, en<br />

drop<strong>in</strong>n sem bara hefur ískr<strong>is</strong>tallamynd<strong>and</strong>i bakteríu<br />

er fros<strong>in</strong>n. Þetta sýnir að ís<strong>in</strong>n hlýtur að vaxa gegnum<br />

loftaugun eða sprungur í yfirhúð<strong>in</strong>ni til að koma<br />

af stað fryst<strong>in</strong>gu í laufblað<strong>in</strong>u (endurprentað eftir<br />

W<strong>is</strong>niewski og Fuller, 1999, með leyfi).

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