Black currant physiology in a changing climate

A. Sønsteby1 and O.M. Heide2
1Bioforsk - Norwegian Institute for Agricultural and Environmental Research, Kapp, Norway; 2Dept. of Ecology
and Natural Resource Management, Norwegian University of Life Sciences, As, Norway.

Abstract
Black currant is a woody plant in which growth and development are intimately
controlled by, and synchronised with seasonal changes in photoperiod and
temperature. Concern over the potential impact of global warming on plant phenology
and yield, led us to initiate relations. An experimental system with single-stemmed
potted plants was developed which allowed a research program to address both
qualitative and quantitative assessment of climatic responses. Growth cessation and
flowering were both induced by short days, with critical photoperiods of
approximately 17 and 16 h, respectively, for most cultivars. Both processes were
advanced and promoted by increasing autumn temperature with an optimum in the
18-21°C region. An exception was cultivars of high-boreal origin, which had an early
growth cessation at low temperature. Unexpectedly, however, not all plants flowered
after exposure to 10 h photoperiod, and the number of flowers decreased as the
photoperiod was reduced from the near-critical length of 15 h. This was due to
premature dormancy induced by an abrupt change to photoperiods well below the
critical level. Field experiments revealed that cultivars of varying geographic origin,
exhibited a typical latitudinal cline in their photoperiodically controlled timing of
growth and flowering responses. Breaking of bud dormancy and promotion of flower
bud development required chilling at -5°C for 14 weeks or more for optimal
responses. However, while chilling at -10°C for 8 weeks resulted in dormancy release,
continued chilling to 16 weeks inhibited bud break completely. We therefore propose
that excessive chilling induces secondary bud dormancy in black currant. The
observed high chilling requirements of black currants concur with the reported
vulnerability of this crop to declining winter chill in the wake of the ongoing global
warming. Furthermore, such conditions also induce a particularly deep bud dormancy
state that further increases the chilling need.

Keywords: black currant, chilling, dormancy, flowering, growth cessation, photoperiod,

Ribes nigrum, temperature

INTRODUCTION
Black currant (Ribes nigrum L.) is an important soft fruit crop for the cold and
temperate regions. As first reported by Nasr and Wareing (1958, 1961a, b) and later
confirmed by Tinklin et al. (1970), initiation of floral primordial requires short day (SD)
conditions in this species. Under field conditions floral initials are formed in late summer
immediately after extension growth has slowed down (Nasr and Wareing, 1961a), and then
develop to anthesis in the following spring. Exposure to 16 short days was sufficient to
induce flowering in the cultivar ‘Baldwin’, while 8 days were insufficient (Nasr and Wareing,
1961b). The critical daylength for floral induction was found to be relatively long,
approximately 16 h in ‘Wellington XXX’ (Tinklin et al., 1970). At this daylength, however, at
least 10 weeks of exposure was required for floral initiation. It was further found that young
shoots with fewer than 12-15 nodes (leaves) have not attained “ripeness to flower” and are
thus unable to respond to SD induction (Tinklin et al., 1970).
Since photoperiod is a function of both season and latitude, the time of floral initiation
will vary with latitude. Thus, collated data for a range of locations in Europe showed a
general correlation between increase in latitude and lateness of initiation (see Tinklin et al.,
1970). The range varied from mid-May-early June at Bologna, Italy (44°30’N) to late June-

  Acta Hortic. 1133. ISHS 2016. DOI 10.17660/ActaHortic.2016.1133.24 159

Proc. XI Int. Rubus and Ribes Symp.
  Eds.: G.E. Fernandez and K.E. Hummer
early July in southern England (51°N) and early August in southern Scandinavia (60°N) to
late September at the arctic circle in Rovaniemi, Finland (cf. Hå rdh and Wallden, 1965;
Vestrheim, 1972). Although this trend may be interpreted as a straightforward latitudinal
effect of photoperiod, it was interpreted by Tinklin et al. (1970) as a latitudinal effect of
temperature on shoot growth and hence, the attainment of ripeness to flower. The reasoning
was, that at lower latitudes, where growth starts early and the prevailing daylength never
exceeds the critical level for flowering, initiation will take place as soon as the shoot has
produced the minimum number of nodes. At higher latitudes, where growth starts later and
the growing temperature is lower, the critical shoot size is reached progressively later, and
therefore, initiation will not take place until late in the season when both the critical number
of leaves and the critical day length are attained.
However, besides its effect on shoot growth, temperature is also an important
controlling factor for flowering in many plants, either directly, or by interaction with
photoperiod (Thomas and Vince-Prue, 1997), but this has been little studied in black
currant. Tinklin et al. (1970) subjected plants of ‘Wellington XXX’ to photoperiods of 12, 14
and 16 h combined with night/day temperatures of 12/17 and 22/27°C (12/12 h). The
results demonstrated that while photoperiod was of overriding importance, temperature
was an important modifying factor, especially under near-critical photoperiods. At 14 h
photoperiod where the effect was largest, a rise in temperature of 10°C resulted in a delay in
the formation of normal flowers by 2 to 6 weeks.
Temperature is also of fundamental importance for the regulation of dormancy in
plants. In the great majority of temperate zone woody plants, growth cessation and bud
dormancy are induced by short photoperiods in the autumn, while release from dormancy
requires exposure to low (chilling) temperature for extended periods (for review, see Vegis,
1964). With the predicted and ongoing global warming (Sunley et al., 2006), there is
increasing concern in parts of Europe that the performance of some soft fruit species, and
particularly black currant, may be adversely affected by mild winters with insufficient
winter chill, leading to uneven and poor bud development in spring (Rose and Cameron,
2009; Jones et al., 2013). The adverse effect of high temperature on flower initiation in black
currant reported by Tinklin et al. (1970) also raises concern about the potential adverse
effects of elevated autumn temperature for floral initiation in this species. This concern is
accentuated by an increasing interest in production of black currants for fresh consumption,
which would demand abundant flowering and large inflorescences that would facilitate hand
harvesting. Also, the early investigations by Nasr and Wareing (1961a, b) and Tinklin et al.
(1970) revealed a coincidence between a decline in extension growth in autumn and floral
initiation. Consistently, floral initiation was preceded by reduced extension growth while
complete growth cessation took place later. However, the causal relationship between the
two developmental processes is not clear.
Faced with these uncertainties and the incomplete knowledge about the physiological
effects of photoperiod and temperature on growth and flowering of black currant, together
with the related concern over the potential impact of global warming on the commercial
cropping of this species, we have initiated a research program to address some of these
relations. Here, we present the results of a range of experiments in black currant cultivars of
contrasting geographical origin.

MATERIALS AND METHODS

Plant material and cultivation

Bushes of 19 black currant cultivars from a field repository were used to monitor
seasonal time-courses of growth cessation and flower bud initiation during two growing
seasons. The repository was located in the central part of South Norway (60°40’N; 10°52’E;
250 m a.s.l.) (for details, see Sønsteby and Heide, 2013). Controlled environment
experiments were also conducted with a range of commercially grown Scandinavian and
Scottish black currant cultivars as well as the high-boreal Russian cultivars ‘Imandra’ and
‘Murmanschanka’, originating from the Kola Peninsula. For these experiments, a system with

160
single-stemmed plants propagated from semi-softwood cuttings was developed as described
by Sønsteby and Heide (2011). The use of plants with a single stem allowed the assessment
of both qualitative and quantitative developmental responses. A coarse-textured sphagnum
peat growth medium with a pH of 5.8 and fertilised with OsmocoteTM controlled-release
fertiliser, was used throughout, as described by Sønsteby and Heide (2011). The plants were
raised in a greenhouse at 20°C in 24-h long days (LD), usually until they had produced 20 or
more nodes. They were then moved into the As phytotron and exposed to different
combinations of temperature and photoperiod as described by Sønsteby and Heide (2011),
Sønsteby et al. (2012), and Heide and Sønsteby (2011, 2012, 2015). Following these
treatments, the plants were stored in the dark at 0±1°C for 12-15 weeks for breaking of bud
dormancy, and finally forced in a greenhouse at 18-20°C and LD conditions to monitor bud
break and flowering performance.
For determination of the chilling requirements of dormant buds, intact plants and
detached shoots of a range of cultivars of widely different origin were stored at
temperatures ranging from 10 to -10°C for 4-21 weeks and forced as single node cuttings,
entire shoots, or intact plants at 20°C and 24 h photoperiod (Sønsteby and Heide, 2014). The
influence of autumn temperature on bud dormancy state was studied by exposing intact
plants of three cultivars to temperatures of 9, 15, and 21°C for 6 weeks during SD dormancy
induction, followed by subsequent chilling at 0°C for 5, 10, or 15 weeks and forcing at 20°C
and LD conditions (Sønsteby and Heide, 2014).

Experimental design, data collection and analysis

All experiments were fully factorial, with a split-plot design, and replicated in three
randomized blocks, each containing 3 to 5 plants of each cultivar for each treatment studied.
In the different studies, extension growth was monitored, by weekly measurements of plant
heights, and the final number of nodes was recorded in each plant at the end of the
treatment period. During forcing, dates of bud burst and anthesis (first open flower) were
recorded by daily observations. The number and position of flowering nodes together with
the total number of flowers at each node were also recorded in each plant at the end of the
flowering period.
The chilling experiments were replicated in three blocks, each containing five single
node cuttings, or three entire shoots or potted plants of each cultivar in each treatment.
During forcing, bud break was observed three times weekly and scored according to the
stages shown in Sønsteby and Heide (2014). Stage 2 was considered as bud break. In
addition, when the observations were terminated, the number and distribution of sprouted
buds were recorded along the entire shoot length of the entire shoots and intact plants, as
well as the number of flowers at each node.
All data were subjected to analysis of variance (ANOVA) by standard procedures using
a MiniTab® Statistical Software program package (Release 16; Minitab Inc., State College, PA,
USA). Percentage values were always subjected an arc sin transformation before
performance of the ANOVA.

RESULTS AND DISCUSSION

Seasonal timing of growth cessation and floral initiation

When growth and development of field grown bushes of 19 black currant cultivars of
different latitudinal origin were studied in a cultivar field repository at 60°40’N (Sønsteby
and Heide, 2013), they exhibited widely different seasonal timing of growth cessation and
floral initiation (Figures 1 and 2). On a whole, the 19 cultivars constituted a typical
latitudinal cline in their photoperiodically controlled timing of growth and flowering
responses. High latitude cultivars originating from crosses and selections of local, wild black
currant populations from the Kola Peninsula and Swedish Lapland were particularly early
and ceased growing and initiated floral primordia already by mid-June. This was approx. 5-6
weeks earlier than any of the other cultivars from lower latitudes. The timing of growth
cessation was closely related to the timing of floral initiation. In general, shoot growth

161
started to slow down approximately 2 weeks before floral primordia became visible in the
dissected buds, while the complete cessation of shoot growth usually followed 1 week later.
With their unique photoperiodic responses, the high-latitude Russian cultivars ‘Imandra’
and ‘Murmanschanka’ represent valuable additions to the limited diversity of the available
black currant gene-pool, and may be of particular use for breeding of cultivars adapted to
the sub-Arctic environment.

Figure 1. Time-courses of shoot elongation growth in 15 black currant cultivars grown in
the field at the Bioforsk Experimental Centre Apelsvoll, Norway (60°40’N).
Seasonal changes in the natural photoperiod are indicated. Each datum point
represents the mean of nine shoots from nine different bushes (after Sønsteby
and Heide, 2013).

Figure 2. Time-courses of flower bud initiation and differentiation in 19 black currant
cultivars grown in the field at the Bioforsk Experimental Centre Apelsvoll, Norway
(60°40’N) during the growing seasons 2011. Each datum point represents the
mean of five buds from five different bushes (after Sønsteby and Heide, 2013).

162
 When six of the same cultivars were grown in phytotron compartments under
constant temperatures of 9, 15 and 21°C and natural day-length conditions at 59°40’N
during the entire growing season, the decline and cessation of growth were advanced by
increasing temperature in all cultivars except the high-latitude cultivar ‘Murmanschanka’. In
‘Ojebyn’, ‘Kristin’, ‘Ben Tron’, ‘Ben Hope’ and ‘Narve Viking’ growth started to level-off at 21°C
by July 20, followed by a complete cessation of growth 2 weeks later (Figure 3). At
temperatures of 15 and 9°C, both stages were delayed by approximately 2 and 4 weeks,
respectively. However, an opposite trend was observed in ‘Murmanschanka’. In this cultivar,
growth started to slow down almost immediately at 9°C with complete growth cessation by
June 15, when the plants had reached a height of only 10 cm. At higher temperatures, growth
continued for approximately a further 2 or 4 weeks, at 15 and 21°C, respectively. However,
even at 21°C, which supported the longest duration of growth, ‘Murmanschanka’ had
completely ceased growing by July 20, at which time the growth of the other five cultivars
had just started to level-off (Figure 3).

Figure 3. Time-courses of cumulative shoot elongation in six black currant cultivars grown
in phytotron daylight compartments under natural photoperiodic conditions at
As, Norway (59°40’N) and constant temperatures of 9, 15, or 21°C, as indicated.
All values are the means of three replicates, each consisting of three plants of each
cultivar (after Sønsteby et al., 2012).

Effects of elevated autumn temperature

When plants of five cultivars were exposed to natural photoperiods and constant
temperatures (9, 12, 18, and 24°C) in phytotron compartments at 59°40’N, growth started to
slow down from August 11, and stopped within 2-3 weeks. Unexpectedly, both growth
cessation and floral initiation were promoted and advanced by increasing temperature. The
number and percentage of flowering nodes increased with increasing temperature in all

163
cultivars, while the corresponding number and proportion of vegetative nodes decreased
(Figure 4). The number of flowers per plant also increased several-fold when temperature
was increased from 9 to 18°C. This is at variance with the results of Tinklin et al. (1970),
who found that under 14 h photoperiod, a 10°C rise in temperature delayed floral initiation
by 2-6 weeks in ‘Wellington XXX’. However, the results were consistently confirmed in
further experiments. It is also a general experience that relatively high temperatures
enhance the effect of SD on the induction of flowering in a range of SD plants (Thomas and
Vince-Prue, 1997; Sønsteby and Heide, 2008). Our results also confirm the close connection
between growth cessation and floral initiation reported previously (Nasr and Wareing,
1961a; Tinklin et al., 1970). Apparently, the two effects are parallel responses to the same
environmental signal (Sønsteby and Heide, 2013).

Figure 4. Appearance of black currant cultivars ‘Kristin’ (A) and ‘Narve Viking’ (B) plants
during flowering in Spring as affected by exposure to varying temperatures as
indicated in the previous Autumn. The lines of demarcation represent the main
position of the lowermost flowering node at each temperature (after Sønsteby
and Heide, 2011).
When actively growing, single-stemmed plants of three black currant cultivars with
15-16 nodes were exposed to photoperiods of 10, 14, 15, 16, 17, 18, 20, and 24 h at 18°C for
8 weeks, growth cessation was induced in all cultivars by short photoperiods with a critical
photoperiod of approximately 16 h. The shorter the photoperiod, the more rapid was the

164
cessation of growth. The appearance of the plants after 6 weeks at various photoperiodic
treatments is shown in Figure 5. An unexpected result was that while all plants flowered
after exposure to 14, 15, or 16 h photoperiod, only 67% (‘Ben Tron’) to 89% (‘Kristin’) of the
plants flowered in 10 h treatment. Also the number of flowers per plant decreased several-
fold in all cultivars as photoperiod was reduced from 15 to 10 h. In parallel with this, the
number of flowers per inflorescence was also nearly halved (see Heide and Sønsteby, 2011).
This was unexpected as photoperiods of 8 h were highly effective in earlier investigations
with other cultivars (Nasr and Wareing, 1961b; Tinklin et al., 1970). The latter authors also
found that 12-h photoperiod was more effective than 14- and 16-h photoperiods. It is also a
definition of SD plants in general, that they flower earlier and more profusely with
decreasing length of day (Thomas and Vince-Prue, 1997). The flowering result was even
more surprising as growth cessation was enhanced by decreasing photoperiod and was
optimal in 10-h photoperiod (Figure 5). The result has been confirmed in further
experiments which show that an abrupt change to photoperiods well below the critical
length induces premature dormancy that arrests floral initiation at an early stage.
Accordingly, the optimum photoperiod for flowering in black currant must be short enough
to allow flower initiation and still long enough to prevent premature dormancy (Heide and
Sønsteby, 2015). Therefore, under naturally decreasing autumn photoperiods, both
processes proceed harmoniously in parallel, and flowering will be optimal.

A 10h 14h 15h 16h 17h 18h 24h B

Figure 5. Appearance of black currant plants after 6 weeks of exposure to different
photoperiods at 18°C. A) ‘Ben Tron’ plants exposed to seven different
photoperiods as indicated; and B) plants of all three cultivars (i.e., one replicate)
exposed to a 10 h (left) and a 24 h photoperiod (right), respectively (after Heide
and Sønsteby, 2011).
In addition to the direct negative effect of low growth temperature on floral initiation,
low temperature during the growing season also had an indirect effect due to reduced and
delayed shoot growth. Thus, under natural day-length conditions during late summer and
early autumn, floral initiation in all cultivars except ‘Murmanschanka’ was progressively
delayed by decreasing temperature, and this was accompanied by parallel reductions of
flowering in the following season (data not shown). This growth-retarding effect of low
temperature delayed, or in some cases even prevented, the plants from reaching the stage of
“ripeness-to-flower” before the treatments were terminated by September 21. Thus, the
cultivars ‘Ojebyn’ and ‘Murmanschanka’, which grew poorly at 9°C, did not flower at all, or
flowered only marginally, at this temperature.
The results also demonstrate again, that the cultivar ‘Murmanschanka’ differed

165
fundamentally from the other cultivars in its environmental responses. While in all the other
cultivars, the slowing down and cessation of growth were advanced by high temperature in
parallel with floral initiation (cf. Sønsteby and Heide, 2011), the opposite trend was found in
‘Murmanschanka’. With its origin at the Kola Peninsula, it appears that this cultivar, like high-
latitude provenances of several other woody plants in general (Heide, 1974; Hå bjørg, 1978),
requires almost continuous light for maintenance of growth. While most modern black
currant cultivars grown commercially in western Europe, vary little in their environmental
responses and seem to represent a common gene-pool (see discussion by Sønsteby and
Heide, 2011; Heide and Sønsteby, 2011), the present results with ‘Murmanschanka’ suggest
a distinct genetic constitution of this and related cultivars of high-boreal origin.

Chilling requirements for breaking of bud dormancy

Chilling requirements for bud dormancy release and subsequent flower development
in contrasting black currant cultivars were assayed at temperatures ranging from +10 to
-10°C using single node cuttings, entire detached shoots, and intact plants (Sønsteby and
Heide, 2014). While single node cuttings underestimated the chilling requirements of intact
plants, severed shoots and intact plants produced similar results. For single node cuttings,
-5°C was the optimal temperature for bud dormancy release. The proportion of breaking
buds increased when storage was extended from 4 to 8 and 12 weeks and then, in most
cases, remained even with further extension to 16 weeks. This was accompanied by a
parallel reduction in the number of days to bud burst (Figure 6). When forcing of entire cut
shoots, chilling at -5°C for 14 weeks or more were optimal for breaking of bud dormancy and
promotion of flower development in most cultivars, flower development usually having
greater chilling needs than bud break itself. All the studied black currant cultivars had
generally low optimum temperatures for breaking of bud dormancy, temperatures of -5 and
-10°C being optimal (Figure 6). However, marked differences in the chilling requirements
were observed between cultivars as illustrated for the cultivars ‘Ben Tron’ and ‘Narve Viking’
in Figure 7. The observed low temperature chilling requirements of black currant buds
concurs with the results of Jones et al. (2013), who found that temperatures of 0°C and
below were optimal for the wide range of cultivars used by them. This is below the +5-0°C
range, which is considered the general temperature optimum for bud dormancy release in
temperate woody plants (Vegis, 1964). This low temperature optimum may explain why
black currant has proved particularly vulnerable to the recent declines in winter chilling due
to global warming (cf. Jones et al., 2013; Atkinson et al., 2013).
Within certain limits, extension of the chilling period compensated for non-optimal
chilling temperature. However, while exposure to -10°C for 8 weeks caused bud dormancy
release, continued chilling for another 8 weeks inhibited bud break completely (Figure 6).
We propose that excessive chilling induces secondary bud dormancy, a principle that is well
established in seed dormancy where it is induced in non-dormant seeds in response to
temperature and light conditions that are unfavourable for seedling growth (Bewley and
Black, 1994). The principle of secondary dormancy has, to our knowledge, not been
recognized in buds of any species before, although an over-chill response was reported by
Jones et al. (2013). Analogous to the situation in seeds, we propose a mechanism whereby
dormancy-released buds are induced to become dormant again (secondary dormancy) if
they remain in low temperature conditions that are unfavourable and even fatal for growing
buds. This would be an important survival mechanism that might have evolved by natural
selection in plants living in seasonally cold and variable environments. In seeds with
secondary dormancy, the dormant state is particularly hard to break, and it would therefore
be interesting to study the chilling requirements for breaking of secondary dormancy in
black currant buds.
It was also found that elevated temperature during SD dormancy induction in autumn
results in a particularly deep state of bud dormancy. When intact plants of three black
currant cultivars were exposed for six weeks to temperatures of 9, 15, or 21°C and natural
day-length at 59°40’N from September 10, and subsequently chilled at 0°C for 5, 10, or 15
weeks and forced at 20°C in LD, it was found that elevated autumn temperature significantly

166
increased the chilling need for breaking of bud dormancy (Figure 8). Both bud burst and
flowering were affected. Modification of the bud dormancy state by temperature during
dormancy induction in autumn is a well-established phenomenon, not only in black currant
(Sønsteby and Heide, 2011; Sønsteby et al., 2012), but in a wide range of temperate trees
and shrubs (Må ge, 1975; Heide, 2003; Palonen, 2006). Elevated autumn temperature
renders the buds in a particularly deep state of dormancy that is manifest as increased
chilling requirements and delayed bud break in spring. This will further increase the large
chilling needs of dormant black currant buds.

30 10°C 100
'Ben Tron'
5°C 90
25 0°C 80

Percent bud break

Days to bud break

-5°C
20 -10°C 70
60
15 50
40
10 30
5 20
10
0 0
30 4 8 12 16 10°C 100 4 8 12 16
'Hedda'
Time (weeks of chilling) Time (weeks of chilling)
5°C 90
25 0°C 80
Days to bud break

Percent bud break

-5°C
20 70
-10°C
60
15 50
40
10
30
5 20
10
0 0
30 4 8 12 16 4 8 12 16
10°C
Time (weeks of chilling) 100 Time (weeks of chilling)
'Imandra' 5°C 90
25 0°C 80
Percent bud break
Days to bud break

-5°C
20 -10°C 70
60
15 50
40
10 30
20
5
10
0 0
4 8 12 16 4 8 12 16
30 10°C
Time (weeks of chilling) 100 Time (weeks of chilling)
'Narve
5°C 90
25 Viking' 0°C 80
Percent bud break
Days to bud break

-5°C
20 -10°C 70
60
15 50
40
10
30
5 20
10
0 0
4 8 12 16 4 8 12 16
Time (weeks of chilling) Time (weeks of chilling)
Figure 6. Bud break in single node cuttings of four black currant cultivars as influenced by
increasing length of chilling at five different temperatures as indicated. Values are
the means ±SE of three replicates, each with five cuttings of each cultivar in each
treatment (after Sønsteby and Heide, 2014).

167
 ‘Ben Tron’ ‘Narve Viking’

Figure 7. Appearance of shoots of ‘Ben Tron’ and ‘Narve Viking’ black currants chilled for
14 weeks at temperatures of 10, 5, 0, -5, and -10°C (from left to right) and forced
for 21 days at 20°C in 24 h photoperiod. Note the effect of low chilling
temperature on the advancement of bud break towards the base of the shoots of
‘Narve Viking’ (after Sønsteby and Heide, 2014).

‘Imandra’ 15°C 21°C ‘Ben Tron’ 15°C ‘Narve Viking’

9°C 9°C

15°C
21°C

9°C 21°C

Figure 8. Effect of temperatures as indicated during autumn dormancy induction on the
advancement and distribution of spring bud break in plants of three black currant
cultivars. Photo taken after 15 weeks of chilling at 0°C and subsequent forcing in
LD at 20°C for 14 days (after Sønsteby and Heide, 2014).

168
CONCLUSIONS
Overall, our results demonstrate a typical latitudinal cline in the photoperiodically
controlled growth and flowering responses of black currant cultivars of varying geographic
origin. Furthermore, the very early cessation of growth, even before mid-summer, as found
in the studied high-latitude black currant cultivars, suggests that these plants of sub-Arctic
origin require almost continuous daylight (midnight sun conditions) to maintain shoot
growth. Clearly, the high-latitude Russian cultivars are genetically distant from the West-
European commercial black currant cultivars and are uniquely adapted to the sub-Arctic
environment, and may therefore represent valuable additions to the limited diversity of the
available black currant gene-pool.
The present results also show that summer temperature affects flowering in black
currant both directly, by promoting and enhancing the SD-induced floral initiation during
late summer and autumn, and indirectly, by the enhancement of growth, resulting in an
advancement of the transition from the juvenile state, and the production of long shoots
with many nodes as potential sites for flowering. Also, the relatively high temperature
optimum for floral initiation shows that, contrary to expectations, there is little reason to
worry about adverse effects of elevated autumn temperature on flower formation in this
crop. Especially in the cool Nordic environment where autumn temperatures obviously are
sub-optimal for SD floral induction, an increase in autumn temperature would actually be
beneficial for floral initiation of the black currant. Likewise, elevated autumn temperature
was found to promote and advance SD induced growth cessation and dormancy
development, and in fact, induced a particularly deep state of winter bud dormancy that is
manifest in prolonged rest and increased chilling requirements for dormancy release.
Experiments with artificial chilling of severed shoots and intact plants have also
demonstrated that temperatures as low as -5°C and below for 14 weeks or more are optimal
for breaking of bud dormancy and promotion of flower development in most black currant
cultivars. The observed high chilling requirement concurs with the reported vulnerability of
this crop to declining winter chill in many parts of the world in the wake of the ongoing
global warming. The finding that elevated autumn temperature increases the depth of
winter dormancy may have important adaptive implications under a scenario of climatic
warming, as it may counterbalance the tendency to premature bud burst in mild winters and
hence, reduce the risk of late frost damage. However, it will further increase the chilling
demand and further accentuate the reported problem of inadequate winter chill in black
currant in regions with mild winters. Breeding of new cultivars with modified chilling
requirements will thus have to compromise between risks of late frost and inadequate
chilling.
Excessive chilling at freezing temperatures inhibited dormancy release. We propose
that this is due to induction of secondary dormancy, a principle that is well established in
seeds, but has to our knowledge, not been recognized for bud dormancy before. Clearly, this
is a phenomenon that needs to be further studied in this and other temperate woody plants
in the present scenario of global warming.

Literature cited
Atkinson, C.J., Brennan, R.M., and Jones, H.G. (2013). Declining chilling and its impact on temperate perennial
crops. Environ. Exp. Bot. 91, 48–62 http://dx.doi.org/10.1016/j.envexpbot.2013.02.004.
Bewley, D.J., and Black, M. (1994). Seeds. Physiology of Development and Germination, 2nd edn (New York, USA:
Plenum).
Hå bjørg, A. (1978). Photoperiodic ecotypes in Scandinavian trees and shrubs. Meld. Norges Landbrukshøgsk. 57
(33), 1–20.
Hå rdh, J.E., and Wallden, J. (1965). Flower formation and fruit growth in black currants. J. Agr. Sci. Finl. 37, 61–75.
Heide, O.M. (1974). Growth and dormancy in Norway spruce ecotypes (Picea abies). I. Interaction of photoperiod
and temperature. Physiol. Plant. 30 (1), 1–12 http://dx.doi.org/10.1111/j.1399-3054.1974.tb04983.x.
Heide, O.M. (2003). High autumn temperature delays spring bud burst in boreal trees, counterbalancing the effect
of climatic warming. Tree Physiol. 23 (13), 931–936 http://dx.doi.org/10.1093/treephys/23.13.931. PubMed

169
Heide, O.M., and Sønsteby, A. (2011). Critical photoperiod for short day induction of flowering in black currant
(Ribes nigrum L.). J. Hortic. Sci. Biotechnol. 86, 128–134.
Heide, O.M., and Sønsteby, A. (2012). Floral initiation in black currant cultivars (Ribes nigrum L.): effects of plant
size, photoperiod, temperature, and duration of short day exposure. Sci. Hortic. (Amsterdam) 138, 64–72
http://dx.doi.org/10.1016/j.scienta.2012.02.008.
Heide, O.M., and Sønsteby, A. (2015). Simultaneous dormancy induction interferes with short day floral induction
in black currant (Ribes nigrum L.). Sci. Hortic. (Amsterdam) 185, 228–232 http://dx.doi.org/10.1016/
j.scienta.2015.02.002.
Jones, H.G., Hillis, R.M., Gordon, S.L., and Brennan, R.M. (2013). An approach to the determination of winter chill
requirements for different Ribes cultivars. Plant Biol (Stuttg) 15 (Suppl 1), 18–27 http://dx.doi.org/10.1111/
j.1438-8677.2012.00590.x. PubMed
Må ge, F. (1975). Dormancy in buds of red raspberries. Meld. Norges Landbrukshøgsk. 54 (21), 1–24.
Nasr, T.A.A., and Wareing, P.F. (1958). Photoperiodic induction of flowering in black currant. Nature 182 (4630),
269 http://dx.doi.org/10.1038/182269a0.
Nasr, T.A.A., and Wareing, P.F. (1961a). Studies on flower initiation in black currant I. Some internal factors
affecting flowering. J. Hortic. Sci. 36, 1–10.
Nasr, T.A.A., and Wareing, P.F. (1961b). Studies on flower initiation in black currant II. Photoperiodic induction of
flowering. J. Hortic. Sci. 36, 11–17.
Palonen, P. (2006). Vegetative growth, cold acclimation, and dormancy as affected by temperature and
photoperiod in six red raspberry (Rubus idaeus L.) cultivars. Eur. J. Hortic. Sci. 71, 1–6.
Rose, G.A., and Cameron, R.W. (2009). Chill unit models for black currant (Ribes nigrum L.) cultivars ‘Ben Gairn’,
‘Ben Hope’ and ‘Ben Tirran’. Sci. Hortic. (Amsterdam) 122 (4), 654–657 http://dx.doi.org/10.1016/j.scienta.
2009.06.029.
Sønsteby, A., and Heide, O.M. (2008). Environmental control of growth and flowering of Rubus idaeus L. cv. Glen
Ample. Sci. Hort. 117 (3), 249–256 http://dx.doi.org/10.1016/j.scienta.2008.05.003.
Sønsteby, A., and Heide, O.M. (2011). Elevated autumn temperature promotes growth cessation and flower
formation in black currant cultivars (Ribes nigrum L.). J. Hortic. Sci. Biotechnol. 86, 120–127.
Sønsteby, A., and Heide, O.M. (2013). Variation in sesonal timing of flower bud initiation in black currant (Ribes
nigrum L.) cultivars of contrasting origin. J. Hortic. Sci. Biotechnol. 88, 403–408.
Sønsteby, A., and Heide, O.M. (2014). Chilling requirements of contrasting black currant (Ribes nigrum L.)
cultivars and the induction of secondary bud dormancy. Sci. Hortic. (Amsterdam) 179, 256–265 http://dx.doi.
org/10.1016/j.scienta.2014.09.038.
Sønsteby, A., Opstad, N., and Heide, O.M. (2012). Effects of summer temperature on growth and flowering in black
currant cultivars (Ribes nigrum L.). J. Hortic. Sci. Biotechnol. 87, 157–164.
Sunley, R.J., Atkinson, C.J., and Jones, H.G. (2006). Chill unit models and recent changes in the occurrence of
winter chill and spring frost in the UK. J. Hortic. Sci. Biotechnol. 81, 949–958.
Thomas, B., and Vince-Prue, D. (1997). Photoperiodism in Plants, 2nd edn (London, UK: Academic Press).
Tinklin, I.G., Wilkinson, E.H., and Schwabe, W.W. (1970). Factors affecting flower initiation in the black currant
Ribes nigrum (L.). J. Hortic. Sci. 45, 275–282.
Vegis, A. (1964). Die Bedeutung von physicalischen und chemischen Aussenfaktoren bei der Induktion und
Beendigung von Ruhezustanden bei Organen und Geveben hö herer Pflanzen. In Encyclopedia of Plant Physiology,
vol. XV/2, W. Ruhland, ed. (Berlin, Germany: Springer-Verlag), p.534–668.
Vestrheim, S. (1972). Flower initiation in black currant in a Southern Norwegian location. Meldinger fra Norges
Landbrukshøgskole 51 (9), 1–5.

170