BOTANY: THE STATE OF THE ART

What Determines a Plant's Cold Hardiness?

John W Einset
Cold-tolerance depends upon a plant's ability to keep water from leaving its cells and
freezing, which severely dehydrates the cells

Banana plantations along Cape Cod, peas and potatoes, survive brief periods
orange groves in the Berkshire Mountains, of frost but are killed when freezing
tropical landscaping in New England! conditions continue for more than about
Unlikely as these images seem, they are four hours. Cold-hardy plants, on the
not entirely out of the realm of possibility, other hand, tolerate extended periods
especially if more can be learned about of freezing, and laboratory tests indicate
the basic mechanisms that govern the that cold hardiness in some of these
tolerance of plants to cold. In fact, goals plants per-
less spectacular than these, yet still
highly significant, are achievable in the
near future because both our
understanding of plant physiology and our
ability to manip- ulate plant cold hardiness
have improved.

WIthout question, resistance to low

tem- peratures is a major factor
determining the geographic distribution
of plant species. So- called chilling-
sensitive plants, such as the tropical
banana and the semitropical avoca- do,
can be severely injured or even killed by
long-term exposure to temperatures
(50 degrees Fahrenheit, for example) that
are well above freezing. By contrast,
chilling-resis- tant plants, such as garden
mits them to survive at temperatures as
low as minus 75 degrees Fahrenheit.

What causes such wide variations in the

sensitivity of plants to cold? As a
conse- quence of natural selection, plants
native to a particular hardiness zone are
adapted to the temperature extremes
that occur in their environment. Remove
them from that envi- ronment, and they
mayor may not survive. For example,
banana plants kept at low, but
nonfreezing, temperatures suffer an
imbal- ance In their metabolism that kills
their cells and causes brown necrotic
streaks to appear on fruits. Or, hardier
plants might be killed by frost that
occurs during their normal period of
vegetative growth. Several cultivars of
rhododendrons and azaleas that are
grown successfully in Georgia and the
Carolinas, for instance, are killed by late
frosts when they are transplanted to
New England only because their tender
vegetative buds initiate growth too early
in spring. Other plants fail to survive
because late-summer frosts kill vegetative
shoots before they become accli- mated
to cold temperatures.

During any given year, a species of tree or

shrub adapted to the north-temperate
envi- ronment alternates between
periods of cold
36 Plant Hardiness
hardiness and Fahrenheit to of the cells into protoplasm,
cold sensrtivity, minus the intercellular metab-
The term spaces, where it,
5 degrees
acclimation too, freezes. The
Fahrenheit, while
(hardening) refers amount of ice in
plants of zones
to the transi- tion the intercel- lular
from a sensitive 5, 4, 3, 2, and 1 spaces increases
to a hardy exhibit rapidly as
condition, while progressively additional water
deacclimation greater cold moves out of the
(dehardening) hardiness. cells. Lef
desig- nates the Obviously, plants unchecked, the
hardy-to-sensitive in all of these loss of water
transition. categories can from cells causes
Obviously, the tolerate some severe
seasonal timing below- freezing dehydration. In
of acclima- tion weather; it is the fact, the most
and deacclimation magnitude of their widely held
is of critical tolerance that explanation of
impor- tance in differentiates frost damage in
determining a them. plants is that
plant's cold death is caused
hardi- ness. The When a plant, directly by the
magnitude and regardless of its advanced state of
duration of the hardiness cellular
acclimated state classification, is dehydration that
are also crucial. In injured by a results when ice
fact, the Arnold killing frost, forms in tissues.
Arboretum's several harmful According to this
hardiness zones processes are explana- tion when
classify woody- involved. One of the concentation
plant species the earliest and of water in cells
according to the most critical falls below a
mag- nitude of processes is the critical "threshold"
the cold tolerance formation of ice value, pro- tein
they exhibit in crystals in the molecules in the
their acclimated spaces between cells' protoplasm
states. Zone 6 their cells. begin to cross-hnk
plants, for Freezing of the with each other,
instance, can water in the forming a sta- ble
withstand intercellular but nonfunctional
minimum spaces causes matrix. In this
temper- atures of water in the per- manently
plus 5 degrees adjacent living altered state of
cells to move out
ohsm slows to a generate high second, the heat
standstill and, mternal contents of (known to physicists
organic solutes as the heat of
since the cells die,
(dissolved com- fusion) given off
the entire plant pounds) during when water on the
dies. acclimation. Finally, surface of a plant
certain plants are freezes, counteracts
Apparently, species
cold tolerant simply the effect of
of plants that
because they can freezing temperatures
survive temperatures
recover from even on water WIthin
lethal to other
the extreme the plant. In
species do so by
dehydra- tion that Massachusetts, for
preventing the
accompanies ice example, cranberry
dehydration caused
formation. Exam- grow-
by ice for- mation.
ples of such species
One way in which
are paper birch
they accomplish this
(Betula papyriieras,
involves
trembling aspen
"supercooling"-the
(Populus tremu-
absence of ice
loides), and several
formation even
willows.
during periods of
freezing In view of all these
temperatures. considerations,
Another way is for what practically can
ice to form in the be done to prevent
intercellular spaces freezing injury in
but for the loss of plants? An obvious
water from cells to strategy is to ensure
be reduced. Often, that plants are
this means of frost well-watered before
prevention involves periods of potential
osmotic alterations frost. By keeping
in the protoplasm their tissues turgid,
of hardy plants. or swollen, one
Halophytes (salt- might be able to
tolerant plants), prevent the extreme
for exam- ple, cellular dehydration
usually are hardier that usually kills
than their non-salt- frosted plants. A
tolerant relatives related treatment is
because the higher to spray tender
osmotic plants with water
concentration of whenever
their protoplasm temperatures are
effectively prevents below freezing. The
water from leaving rationale of this
cells and con- procedure is
tributing to twofold. First, it
extracellular ice maximizes the
crystals. Some other water con- tent of
hardy plants living tissues and,
Plant Hardiness 37
ers routinely use water spnnklers in
their bogs during late-spring frosts to take
advan- tage of the heat of fusion released
when water freezes on the surface of
cranberry plants. The heat released by
100 gallons of water when it freezes is
approximately equivalent to the amount
of heat produced by burning one gallon of
fuel oil. Obviously, the sprink- ler
technology is an important frost-protec-
tion measure. Breeding programs to
mtro- duce hardiness genes into less
hardy plants may also become extremely
important in the next few years.
Other ways of preventing frost injury in
plants are still in the expenmental stages
and therefore are not yet of practical value.
Some,
in fact, are quite controversial.
Researchers at the University of California,
for example, are attempting to utilize
genetically engi- neered strains of bactena
as frost-protection agents. They reason
that some bacterial spe- cies called "ice-
nucleating bacteria," nor- mally
associated with plants, tend to sensi- tize
plants to freezing injury, since individual
bacteria act as "nucleation centers" for
the formation of ice crystals. Other
bacterial spe- cies, by contrast, are
nonnucleating. Dis- place ice-nucleating
bacteria on a plant with nonnucleating
ones, it is argued, and the plant should
be less prone to frost injury.
Unfortunately, it is still too early to
judge whether theory and practice are
compatible

The paper birch (Betula papynfera). In the Arnold Arboretum (left). and the trembling aspen (Populus trernuloides],
In the San FranCISCO MountaInS. Anzona. are cold tolerant sttnply because they can recover from the extreme
dehydration due to the [ormation of Ice In their Intercellular spaces Many other species are cold tolerant because
they prevent dehydrauon of their cells In the first place Photograph of Populus tremuloides by Susan D McKelvey
Both photographs from the ArchIves of the Arnold Arboretum.
38 Plant Hardiness
in this case. Not of the public's
only that, but concerns about
several experts nonnucleating
question the bacteria as frost
wisdom of protectants will
introducing ultimately depend
potentially on the results of
harmful bacteria controlled experi-
into the envi- ments designed to
ronment. They identify possible
identify several adverse
important questions. environmental
Will the engineered effects of this
bacteria cause practice. If none
undesirable plants, are found, then
such as weeds, to significant progress
become frost will have been
tolerant, too? Or, made on one
since the ice- strategy for
nucleat- ing manip- ulating
bacteria normally plant cold
present on plants hardiness. As
may have beneficial usual, improved
but unrecognized scientific
effects on their understanding of
host plants, might a phe- nomenon,
not their such as cold
beneficial effects be hardiness, if it is
abolished? Lastly, properly applied,
is it possible that has the potential
nonnucleating of improv- ing
bacteria could horticultural
affect the weather, technology.
a prospect that
could have
profound John W. Einset, a staff
consequences? member of the Arnold
As is the case Arboretum, IS
whenever basic aSSOCIateprofessor of
biology III Harvard
science is used to
Umversity,
solve practical
problems,
resolution