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New Zealand Journal of Botany, 1996, Vol.

34: 273-278 273

0028-825X/96/3402-0273 $2.50/0 © The Royal Society of New Zealand 1996

Comparison of frost tolerance of Nothofagus solandrivar.

cliffortioides (Hook.f.) Poole and Nothofagus menziesii (HooW.i.)

OSBERT J. SUN (Hook.f.) Oerst. (silver beech) occur naturally at

NZ Forest Research Institute Limited similar latitudes and altitudes in New Zealand.
Private Bag 4800 Nothofagus solandri commonly dominates the for-
ests at high altitudes in the drier inland mountain
Christchurch, New Zealand
regions of the South Island; Nothofagus menziesii
dominates or co-dominates with N. solandri at
timberlines with high rainfall and improved soil
School of Forestry
conditions (Wardle 1984). Sun et al. (1995) relate
University of Canterbury the difference in the present distribution of N.
Private Bag 4800 solandri and N. menziesii to differences between the
Christchurch, New Zealand two species in tolerance to water stress and water-
logging. It is not known, however, whether the at-
tributes of other environmental variables, e.g., low
Abstract Frost tolerance was investigated in both winter temperature and summer frosts, are important
cold-hardened and actively growing seedlings of in differentiating the geographical distribution of the
Nothofagus solandri var. cliffortioides and two species.
Nothofagus menziesii in New Zealand. Seedlings of
Frost causes freezing of plant tissues, resulting in
the two species from the same geographical origins
injury due to intracellular and/or extracellular ice
showed similar tolerances to simulated mid winter
formation. Frost tolerance of plants is considered to
and summer frosts when grown in a common envi-
be a genetically determined physiological trait but
ronment. The maximum frost hardiness (LT50)
its expression is determined by environmental con-
ranged from -9.0 ± 0.8 to -12.1 ± 0.6°C in N.
straints (Sakai & Larcher 1987). Most woody plants
solandri, and from-10.0 ± 0.7 to-12.4 ± 0.1°C in
native to New Zealand show well-defined geographi-
N. menziesii. Both species exhibited provenance
cal limits along various temperature-related gradi-
variations in frost tolerance that corresponded with
ents (Sakai & Wardle 1978). Frost hardiness for N.
variation in long-term means of annual minimum
solandri was reported to range from -3.5 to —13°C
temperature and frost days per annum of the seed
in mid winter, apparently related to the altitude of
origins. It is suggested that low temperature may not
origin (Sakai & Wardle 1978; Sakai et al. 1981;
be a key environmental factor in differentiating the
Greer et al. 1989). Greer et al. (1989) suggest that
presence of the two species at upper timberlines in
the altitudinal variations in timberlines formed by N.
New Zealand.
solandri mainly reflect the frost hardiness limits of
seedlings. To elucidate the role of low temperature
Keywords frost tolerance; geographical distribu- in differentiating the natural distribution of N.
tion; Nothofagus menziesii; Nothofagus solandri var. solandri and N. menziesii, we investigated the frost
cliffortioides; provenance; timberline tolerance of both cold-hardened and actively grow-
ing seedlings of the two species with the same seed
origins and grown at the same conditions.
Nothofagus solandri var. cliffortioides (Hook.f.)
Poole (mountain beech) and Nothofagus menziesii MATERIALS AND METHODS
Cold-hardened seedlings
B95034 Seed ofN. solandri and N. menziesii were collected
Received 14 August 1995; accepted 11 December 1995 from Lake Hauroko, Eglinton Valley, and Cobb
274 New Zealand Journal of Botany, 1996, Vol. 34

River, in the South Island, and germinated in a glass- by a 100 watt heating element in the base of each
house at the University of Canterbury, Christchurch, frost trolley. The soil surface was also well insulated.
in spring 1990. At each site, collection was made on Following the completion of all frost treatments,
eight maternal trees with heavy seed crops. The seedlings were transported back to Christchurch and
minimum distance between trees used to provide placed into a glasshouse with temperatures set for
seed was 50 m. Table 1 lists the geographical loca- 15°C by day and 10°C at night. Frost damage was
tions and long-term means of annual minimum tem- scored visually 4 weeks after treatment on a scale
perature and frost days per annum of the three seed of 0, no damage; 1, approx. 10% leaves damaged;
collection sites (provenances). 2, approx. 30% leaves damaged; 3, approx. 50%
Two weeks after germination, seedlings were leaves damaged (LT50); 4, approx. 80% leaves dam-
transplanted into 1 1 pots (15 cm top diameter x aged; and 5, shoot tissue dead, as described by
12 cm height) containing a mix of peat (60%), soils Menzies et al. (1981). Frost hardiness (LT50) was
of a Nothofagus-dominated forest (20%), ground defined as the temperature which caused 50% foliar
pine bark (10%), coarse river sand (10%), and slow- damage (Ritchie 1991). This was determined by
release fertiliser (2 kg/m3 Osmocote®Plus, Grace- drawing free-hand curves of damage rating versus
Sierra, Heerlen, The Netherlands). All seedlings frost temperatures for previously-matched seedlings
were kept in the glasshouse until the end of March based on size and appearance under the three frost
before being transferred into an open shade house treatments. Analysis of variance (ANOVA) was
to induce cold hardiness. The one-year-old seedlings conducted on the estimates of LT 5 Q.
were transported to Palmerston North in July to
undergo the frost treatment using low temperature Actively growing seedlings
controlled environment (C.E.) facilities (Robertham Seedlings of the two species were lifted from the
et al. 1978) at the HortResearch National Climate floor of a mixed Nothofagus forest in the Eglinton
Laboratory, Palmerston North. Valley in early January 1992, and potted into 11 pots
Ninety seedlings, 15 for each provenance in each containing the same potting mix as described above.
species, were divided into 3 groups matched in size The size and appearance of these seedlings were
and appearance, and placed randomly in the low- similar to the cold-hardened seedlings of the previ-
temperature C.E. rooms with frost temperature set ous experiment but their ages were not known. All
for—5, —10, or—13°C. The C.E. rooms were run on seedlings were placed initially in an open shade
a 6—6-4 programme (Greer & Warrington 1982), i.e., house at the University of Canterbury. More than
6 hours for cooling from 12°C, 6 hours being main- 80% of the seedlings had recovered from transplant-
tained at the minimum temperature, and 4 hours for ing after 4 weeks, and these were transported to the
warming up to 12°C. The actual temperatures dur- HortResearch National Climate Laboratory,
ing frosting were recorded as-5.0 ± 0.8, -10.2 ± 0.2, Palmerston North, at the end of February 1992. In-
and—13.9 ± 1.1°C, respectively. The frosts were sufficient seed was available to provide seedlings
conducted in darkness, with the lights turned on one raised from seed in the glasshouse for this experi-
hour before the warming was completed. During the ment.
frost process, soil temperature was maintained at 5°C Seedlings were divided into 4 groups, with each

Table 1 Geographical locations of provenance and long-term means of frost

days per annum and annual minimum temperature (Tmin) of the sites.

Altitude Annual Tmin Map

Provenance Latitude (m a.s.l.) frost (days) (°C) number
Cobb River 41°08'S 870 79 -6.5 NZMS260 M26
Eglinton Valley 45°06'S 270 58 -6.1 NZMS260 D43
LakeHauroko 46°06'S 150 26 -4.9 NZMS260 C43
Source: New Zealand Meteorological Service (1983).
Sun & Sweet—Frost tolerance in Nothofagus 275

Fig. 1 Frost damage in cold-

hardened seedlings subjected to L. Hauroko -o- Eglinton V. • Cobb R.
different temperatures. Vertical
bars show standard errors of
means, n = 5, LT50 is indicated by N. solandri
the broken line. 4H

• ~ -I _

CD 0-

N. menziesii


1 -

0 -2 -4 -6 -8 -10 -12 -14

Frost temperature (°C)

group containing 12 plants of each species. Seedlings RESULTS

in each group were placed randomly in a low-tem-
perature C.E. room set for a specific frost tempera- Cold-hardened seedlings
ture. The rooms were run on the 6-6-4 programme Frost damage of cold-hardened seedlings increased
with temperatures of-1.0 ± 0.1, -2.5 ± 0.1,-3.9 ± with decreasing temperatures for seedlings from all
0.1,and-5.4±0°C. three provenances in both species (Fig. 1). Seedlings
Upon completion of the frost treatments, all seed- subjected to -5°C frost showed only minor damage
lings were brought back to Christchurch and placed in all provenances of both species, but considerable
in a glasshouse at the University of Canterbury for damage occurred to those subjected to approximately
visual assessment of frost damage. The assessment —14°C. In both species, seedlings from the three
was conducted 4 weeks after the treatment, with the provenances differed clearly in damage rating at
number of plants showing production of new shoots —10°C; the most severe damage occurred in seed-
at the stem base after complete shoot damage being lings of the Lake Hauroko provenance, and the least
recorded 6 weeks after treatment. Differences in frost in the Cobb River provenance.
damage between the two species were compared by Frost hardiness (LT50) was significantly (P <
Student's Mest at each frost temperature. 0.001) affected by provenance. In N. solandri, LT50
276 New Zealand Journal of Botany, 1996, Vol. 34

Fig. 2 Frost damage in actively

—_____==-—®— m growing seedlings. Vertical bars
__-—-— show standard errors of means,
n = 12, LT50 is indicated by the
Damage rating

4- broken line.


1- I "*" N. solandri
"•" N. menziesii
0 -1 -2 -3 -4 -5 -6
Frost temperature (°C)
ranged from -9.0 ± 0.8°C for the Lake Hauroko DISCUSSION
provenance, through-10.2 ± 1.0°C for the Eglinton
Valley provenance, to -12.1 ± 0.6°C for the Cobb Low temperature plays a prominent role in the dis-
River provenance; while in TV. menziesii, it was—10.0 tribution of woody plants. Altitudinal limits for tree
± 0.7, —11.3 ± 0.4 and-12.4 ± 0.1°C, respectively. growth are largely determined by low temperature.
There was no significant difference between the two Although N. solandri forms the highest timberline
species in frost hardiness in all three provenances. in New Zealand, its cold tolerance in both cold-hard-
ened and actively growing seedlings was found to
Actively growing seedlings be similar to that in N. menziesii for one-year-old
Frost at -1.0°C caused minor damage to actively plants when the two species were grown in the same
growing seedlings of both N. solandri and N. environments. Both species showed a level of mid
menziesii, while nearly all plants subjected to frost winter frost hardiness similar to that of Nothofagus
temperatures of-2.5°C and below suffered severe cunninghamii (Read & Hill 1988, 1989; Read &
frost injury (Fig. 2). There was no significant differ- Hope 1989), Nothofagus betuloides and Nothofagus
ence between TV. solandri and N. menziesii in the dombeyi (Alberdi et al. 1985), and Nothofagus nitida
damage rating for seedlings subjected to -1.0°C (Sakai et al. 1981); but greater than Nothofagus
frost. LT50 for actively growing seedlings in the two fusca, Nothofagus truncata, and Nothofagus moorei
species is estimated to be at a temperature of approxi- (Sakai etal. 1981; Read & Hope 1989). Two decidu-
mately -1.8°C. ous subalpine Nothofagus species, Nothofagus
Six weeks after the frost treatment, some seed- antarctica which occurs naturally in Chile, and N.
lings subjected to apparent lethal temperatures gunnii which is endemic to Tasmania, were found
showed production of new shoots at the base of the to be the most cold tolerant species in the genus
main stem, especially in TV. menziesii. The percent- (Sakai et al. 1981; Alberdi et al. 1985). The frost
age of plants showing production of new shoots dif- hardiness in cold-hardened N. solandri seedlings
fered considerably between the two species; one-third obtained in this study is consistent with those re-
ofN. menziesii seedlings produced new shoots after ported in other studies (Sakai & Wardle 1978; Sakai
being frosted at-2.5 and-3.9°C, but only one-sixth et al. 1981; Greer et al. 1989). \nN. menziesii, simi-
of N. solandri seedlings at -2.5°C frost or N. lar winter frost hardiness was also reported for adult
menziesii seedlings at—5.4°C were observed to have trees (Neuner & Bannister 1995).
such new shoots. Nothofagus menziesii seedlings The tests of tolerance to summer frost were con-
usually produced several vigorous new shoots on a ducted using seedlings collected from the floor of a
single plant, whereas N. solandri seedlings normally Nothofagus forest in the Eglinton Valley because of
produced a small, single shoot only. the lack of a sufficient seed source. Consequently the
Sun & Sweet—Frost tolerance in Nothofagus 277

age of seedlings tested was unknown. Neverthless, water stress. Nothofagus solandri has been shown
the actively growing seedlings of the two species to be much more tolerant to water stress than N.
exhibited a frost hardiness of approximately—1.8°C. menziesii (Sun et al. 1995). A combination of high
This is consistent with the results obtained by Greer radiation, low soil temperature, and shallow soil may
et al. (1989) who found that the mid summer frost cause severe water stress to plants growing in sub-
hardiness in N. solandri varied from —1 °C for seed- alpine environments (Schulze & Hall 1982).
lings collected from 460 ma.s.l. atGlentui, to-3.5°C
for those collected from 1100 m a.s.l. on the lower
slopes of Mt Cockayne in the Craigieburn Range.
Provenance variation in frost tolerance was
clearly displayed in both TV. solandri and N. ACKNOWLEDGMENTS
menziesii, with the variation corresponding appar- We are grateful to E. A. Halligan and I. Warrington for
ently to variations in long-term means of annual their kind support in use of the low-temperature facili-
minimum temperature and frost days per annum ties at the HortResearch National Climate Laboratory,
among provenances even when seedlings of differ- Palmerston North. We thank D. H. Greer for his techni-
ent seed origins were grown in a common environ- cal support and review of the manuscript. This study was
partially supported by a research grant from the Robert
ment. This suggests that frost hardiness in the two C. Bruce Trust. O. J. Sun was a recipient of a T. W. Adams
species is more related to the environmental condi- Scholarship and BNZ Scholarship for Forestry Research.
tions of their natural habitats than to the genetic dif-
ference between them. It may be postulated that low
temperature has not acted as a prominent selection
pressure to differentiate the present distribution of
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