Professional Documents
Culture Documents
TMP 8 C64
TMP 8 C64
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
2-\
c
• ~ -I _
CD 0-
N. menziesii
2-
1 -
0
0 -2 -4 -6 -8 -10 -12 -14
4- broken line.
/
3-
2-
1- I "*" N. solandri
"•" N. menziesii
n-
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
N. solandri and N. menziesii. REFERENCES
Frost survival capacity depends primarily on the Alberdi, M.; Romero, M.; Rios, D.; Wenzel, H. 1985:
specific frost tolerance of a plant. However, the Altitudinal gradients of seasonal frost resistance
importance of recovery from frost injuries has also in Nothofagus communities of southern Chile.
been stressed by Sakai & Larcher (1987) who stated Oecologia plantarum 6: 21—30.
that the chance of frost survival of a plant in a given Greer, D. H.; Warrington, I. J. 1982: Effect of photope-
environment consisted of mitigation or exclusion of riod, night temperature, and frost incidence on
the freezing risk, frost tolerance, and recovery after development of frost hardiness in Pinus radiata
frost damage. During a summer frost, the basal buds D. Don. Australian journal of plant physiology 9:
of a plant could remain undamaged partly due to 333-342.
insulation provided by litter on the forest floor, and Greer, D. H.; Wardle, P.; Buxton, R. P. 1989: Seasonal
partly due to their relatively high resistance to low frost hardiness of Nothofagus solandri seedlings
temperature compared with leaves and shoots higher from two altitudinally diverse sites in Canter-
above (Larcher & Bauer 1981). Regrowth is thus bury, New Zealand. New Zealand journal of
botany 27: 299-304.
possible in species which are capable of producing
new shoots at their bases after suffering from severe Larcher, W.; Bauer, H. 1981: Ecological significance of
dieback. Although N. solandri and N. menziesii resistance to low temperature. In: Lange, O. L.;
showed similar frost hardiness in seedlings originat- Nobel, P. S.; Osmond, C. B.; Ziegler, H. ed.
Physiological plant ecology I. Responses to the
ing from the same sites, Nothofagus menziesii may
physical environment. Encyclopedia of plant
be more capable of surviving a summer frost than physiology, new series, Vol. 12A. Berlin
N. solandri, because of its greater ability to produce Heidelberg, Springer-Verlag. Pp. 403-474.
new shoots from the base of the stem. Thus, in ar-
Menzies, M. I.; Holden, D. G.; Green, L. M.; Rook, D. A.
eas with a high frequency of summer frost, Notho-
1981: Seasonal changes in frost tolerance of Pinus
fagus menziesii might have a more extended radiata seedlings raised in different nurseries.
distribution than N. solandri. New Zealandjournal of forestry science 11: 100—
The present study suggests that frost tolerance 111.
may not contribute directly to the differences be- Neuner, G.; Bannister, P. 1995: Frost resistance and sus-
tween N. solandri and N. menziesii in forming up- ceptibility to ice formation during natural harden-
per timberlines in New Zealand. The differentiation ing in relation to leaf anatomy in three evergreen
between the two species in dominance at timberlines tree species from New Zealand. Tree physiology
might be related, at least partially, to the effects of 15: 31\-311.
278 New Zealand Journal of Botany, 1996, Vol. 34
New Zealand Meteorological Service. 1983: Summaries Sakai, A.; Larcher, W. ed, 1987: Frost survival of plants.
of climatological observations to 1980. New Zea- Responses and adaptation of freezing stress. Eco-
land Meteorological Service miscellaneous pub- logical Studies 62. Berlin Heidelberg, Springer-
lication 177. 172p. Verlag. 321 p.
Read, J.; Hill, R. S. 1988: Comparative responses to Sakai, A.; Wardle, P. 1978: Freezing resistance of New
temperature of the major canopy species of Tas- Zealand trees and shrubs. New Zealandjournal of
manian cool temperate rainforest and their eco- ecology 1: 51-61.
logical significance. I. Foliar frost resistance. Sakai, A.; Paton, D. M.; Wardle, P. 1981: Freezing resist-
Australian journal of botany 36: 131-143. ance of trees of the southern temperate zone,
Read, J.; Hill, R. S. 1989: The response of some Austral- especially subalpine species of Australasia. Ecol-
ian temperate rain forest tree species to freezing ogy 62: 563-570.
temperatures and its biogeographical significance. Schulze, E.-D.; Hall, A. E. 1982: Stomatal responses,
Journal of biogeography 16: 21—27. water loss and CO2 assimilation rates of plants in
Read, J.; Hope, G. F. 1989: Foliar frost resistance of contrasting environments. In: Lange, O. L.; Nobel,
some evergreen tropical and extratropical Aus- P. S.; Osmond, C. B.; Ziegler, H. ed. Physiologi-
tralasian Nothofagus species. Australian journal cal plant ecology II. Water relations and carbon
of botany 37: 361-37'3. assimilation. Encyclopedia of plant physiology,
Ritchie, G. A. 1991: Measuring cold hardiness. In: Lassoie, new series, Vol.l2B. Berlin Heidelberg, Springer-
J. P.; Hinckley, T. M. ed. Techniques and ap- Verlag. Pp. 181-230.
proaches in forest tree ecophysiology. Boca Raton, Sun, O. J.; Sweet, G. B.; Whitehead, D.; Buchan, G. D.
CRC Press Inc. Pp. 557-582. 1995: Physiological responses to water stress and
Robertham, R. W.; Lloyd, J. B.; Warrington, I. J. 1978: A waterlogging in Nothofagus species. Tree physi-
controlled environment room for producing ology 15: 629-638.
advective white or black frost conditions. Journal Wardle, J. 1984: The New Zealand beeches. Ecology,
of agricultural engineering research 23: 301— utilisation and management. Christchurch, New
311. Zealand Forest Service. 447p.