Current issue: 56(1)
Under compilation: 56(2)
Trees are particularly susceptible to climate change due to their long lives and slow dispersal. However, trees can adjust the timing of their growing season in response to weather conditions without evolutionary change or long-distance migration. This makes understanding phenological cueing mechanisms a critical task to forecast climate change impacts on forests. Because of slow data accumulation, unconventional and repurposed information is valuable in the study of phenology. Here, I develop and use a framework to interpret what phenological patterns among provenances of a species in a common garden reveal about their leafing cues, and potential climate change responses. Species whose high elevation/latitude provenances leaf first likely have little chilling requirement, or for latitude gradients only, a critical photoperiod cue met relatively early in the season. Species with low latitude/elevation origins leafing first have stronger controls against premature leafing; I argue that these species are likely less phenologically flexible in responding to climate change. Among published studies, the low to high order is predominant among frost-sensitive ring-porous species. Narrow-xylemed species show nearly all possible patterns, sometimes with strong contrasts even within genera for both conifers and angiosperms. Some also show complex patterns, indicating multiple mechanisms at work, and a few are largely undifferentiated across broad latitude gradients, suggesting phenotypic plasticity to a warmer climate. These results provide valuable evidence on which temperate and boreal tree species are most likely to adjust in place to climate change, and provide a framework for interpreting historic or newly-planted common garden studies of phenology.
Two dynamic models predicting the development of frost hardiness of Finnish Scots pine (Pinus sylvestris L.) were tested with frost hardiness data obtained from trees growing in the natural conditions of Finland and from an experiment simulating the predicted climatic warming. The input variables were temperature in the first model, and temperature and night length in the second. The model parameters were fixed on the basis of previous independent studies. The results suggested that the model which included temperature and photoperiod as input variables was more accurate than the model using temperature as the only input variable to predict the development of frost hardiness in different environmental conditions. Further requirements for developing the frost hardiness models are discussed.
Seedlings of Picea abies (L.) H. Karst. full-sib families of contrasting origins were cultivated in a phytotron under different photoperiodic, light-intensity and temperature treatments during their first growth period. The effects of the treatments on juvenile growth traits – whether enhanced or delayed maturation was induces – were observed during the two subsequent growth periods. The following hypotheses were tested: (A) Enhanced maturation can be induced in the first growth period from sowing with (i) a long period of continuous light during active growth (24 weeks vs. 8 weeks); (ii) a shorter night during bud maturation (12 h vs. 16 h); high temperature (25°C vs. 20°C) during (iii) active growth, growth cessation and bud maturation; and during (iv) the latter part of growth cessation and bud maturation only. (B) Delayed maturation can be induced after (i) low light intensity during growth cessation and bud maturation (114 μmol m-2 s-1 vs. 340 μmol m-2 s-1); low temperature (15°C vs. 20°C) during (ii) active growth, growth cessation and bud maturation; and during (iii) the latter part of growth cessation and bud maturation only.
The most dramatic effect was observed after 24 weeks of continuous light during active growth. All traits showed a significantly more mature performance in the second growth period compared with the control. The effect for all but one trait was carried over to the third growth period. This is in accordance with the hypothesis that the activity of apical shoot meristems controls the maturation process. For the other treatments there was only weak or no support for the hypothesis of induction of enhanced or delayed maturation. Strong family effects were observed for all traits. Differential responses of the various latitudinal families were observed, suggesting that family effects must be considered to predict and interpret correctly how plants will respond to environmental effects.
Different approaches to the study of the annual rhythm of forest trees are described and compared by analysing the concepts and theories presented in the literature. The seasonality varying morphological and physiological state of forest trees is referred to as the annual rhythm s. lat., from which the annual ontogenetic rhythm is separated as a distinct type. The dormancy phenomena of the trees are grouped into four categories. Theories concerning the regulation of the annual rhythm are divided into two main types, the most common examples of which are the photoperiod theory and the temperature sum theory. Recent efforts towards a synthetic theory are described.
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Eight Scots pine (Pinus sylvestris L.) seed sources, ranging from 42° to 66° north latitude, were grown under a constant, 16-hour photoperiod in a greenhouse for approximately 6 months. Rates of photosynthesis, as measured by an IRGA, and growth, as measured by increase in height and fresh and dry weight, differed among seed sources at the end of the six-month growing period. Photosynthetic capacity and growth were strongly related to latitude of seed source, and were greatest in the seed sources coming from a parent environment in which maximum photoperiods are about 16 hours.
Photosynthetic efficiency (rate of photosynthesis per gram needle weight) was also strongly related to latitude of seed source, but was lowest in the seedlings which exhibited the greatest growth and photosynthetic capacity. This may have been due to (1) more mutual shading of needles on the larger seedlings and (2) a lesser proportion of juvenile needles on the larger seedlings or (3) biochemical differences in the use of photosynthate in the needles. Seed source and light intensity had an interacting effect on rates of photosynthesis only in seedlings of the two northernmost seed sources.
The PDF includes a summary in Finnish.
Male flowering was studied at the canopy level in 10 silver birch (Betula pendula Roth) stands from 8 localities and 14 downy birch (B. pubescens Ehrh.) stands from 10 localities in Finland in 1963–73. Distribution of cumulative pollen catches was compared to the normal Gaussian distribution. The basis for timing of flowering was the 50% point of the anthesis-fitted normal distribution. To eliminate effects of background pollen, only the central, normally distributed part of the cumulative distribution was used. Development was measured and tested in calendar days, in degree days (> 5°C) and in period units. The count of the parameters began in March 19.
Male flowering in silver birch occurred from late April to late June depending on latitude, and flowering in downy birch took place from early May to early July. The heat sums needed for male flowering varied in downy birch stands latitudinally but there was practically no latitudinal variation in silver birch flowering. The amount of male flowering in stands of the both species were found to have a large annual variation but without any clear periodicity.
The between years pollen catch variation in stands of either birch species did not show any significant latitudinal correlation in contrast to Norway spruce stands. The period unit heat sum gave the most accurate forecast of the timing of flowering for 60% of the silver birch stands and for 78.6% of the downy birch stands. Silver birch seems to have a local inclination for a more fixed flowering date compared to downy birch, which could mean a considerable photoperiodic influence on flowering time of silver birch. The species had different geographical correlations.
Frequent hybridization of the birch species occurs more often in Northern Finland than in more southerly latitudes. The different timing in the flowering causes increasing scatter in flowering times in the north, especially in the case of downy birch. Thus, the change of simultaneous flowering of the species increases northwards due to a more variable climate and higher altitudinal variation. Compared with conifers, the reproduction cycles of the two birch species were found to be well protected from damage by frost.
The timing of the tetrad phase of microsporogenesis in sixteen tree species, belonging to the genera of Abies, Larix, Picea, Pinus, Alnus, Betula, Corylus and Populus, was studied. The tetrad phase of microsporogenesis in conifers and in Populus tremula L. was reached from late March to early June including the yearly and latitudinal variation. The tetrad phase in Betulaceae was reached in late July to mid-August. The microsporogenesis in Betulaceae species differed in ecophysiological terms from the other species studied in that the timing in Betulaceae was rather day-length dependent than heat sum-correlated. In conifers and in Populus the timing of tetrad phase correlated with heat sums accumulated and did not correlate with day length or any kind of thermal threshold. This difference was, however, judged to be associated to seasonal adaptive strategies rather than taxonomic relationships.
The PDF includes a summary in Finnish.