Scientia Horticulturae 224 (2017) 61–67

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Review

Late-blooming in almond: A controversial objective MARK

a,⁎ a b
José M. Alonso Segura , Rafel Socias i Company , Ossama Kodad
a
Unidad de Hortofruticultura, Centro de Investigación y Tecnología Agroalimentaria de Aragón (CITA), Av. Montañana 930, 50059 Zaragoza, Spain
b
Département Arboriculture-Viticulture, École Nationale d’Agriculture de Meknès, Meknès, BP S/40, Morocco

A R T I C L E I N F O A B S T R A C T

Keywords: Blooming time is an essential trait in almond cultivars, mainly because of the occurrence of frosts during bloom
Prunus amygdalus and early fruit development, when the flowers and the young fruits are very sensitive to freezing temperatures.
Almond Traditional almond growing has been mostly restricted to coastal regions where the risk of frosts was minimal or
Blooming time nil, but at present it is shifting to two very different environments: inland regions with high frost incidence and
Chilling requirements
subtropical regions with very low chilling accumulation. Therefore, the chilling and heat requirements for al-
Heat requirements
mond blooming have acquired the highest interest when describing a new cultivar and, as a consequence, these
Breeding
requirements have been analysed in this review, as well as the genetics of blooming time as a basis for defining
late bloom as an objective in breeding programmes or not. Additionally, the breeding efforts directed to obtain
later blooming cultivars have been reviewed, stating that no breeding has been yet undertaken towards earlier
bloom because many traditional cultivars were early blooming. Almond breeding has been highly effective in
obtaining very late blooming cultivars, well adapted to growing regions with high frost risks, but these very late
blooming releases may conversely show adaptation problems in low chilling regions.

1. Introduction
Genetic variation for flowering time has been described in natural
populations of most plant species, including almond. It may represent
different adaptations to more coastal or continental climatic conditions
because almond has been the first temperate fruit species to bloom. For
this reason, almond growing was restricted to regions where the risk of
frost was low or even nil. Despite this restriction, over centuries of al-
mond growing many attempts have been undertaken to shift its culti-
vation from the Mediterranean shores towards inland regions where, in
most years, spring frosts occur during the time of almond blooming or
soon after (Socias i Company et al., 2012), greatly limiting its profit-
ability, thus becoming frequently a marginal crop, only for self-con-
sumption.
Almond production shows great variability in many producing
countries (Gradziel et al., 2017). A special case is found in Spain, as
seen by the extreme variability of the Spanish almond production for
the period 2001–2016 (Fig. 1). Most of the variability is due to the
occurrence of frosts, as especially shown in the year 2004, when in
March there were several intermittently injuring frosts every few days,
affecting practically all growing regions and the successively blooming
cultivars (Kodad and Socias i Company, 2005). The study of two groups
of breeding selections of different blooming season showed that the
earlier blooming suffered heavier damages than those of very late
blooming time, some of them without damages because they bloomed
later than the last frost on March 26. However, the differences in frost
damages within each blooming group showed the existence of sig-
nificant differences in intrinsic resistance of each genotype to frost, as
already observed by Felipe (1988). Consequently, both blooming time
and intrinsic resistance must be considered when evaluating the real
level of resistance of a genotype to frost damage (Kodad et al., 2010).
Similar variability in almond production due to frosts has also been
reported in other countries, such as India, Chile and Argentina (Kumar
et al., 1993; Castro, 2013; Iannamico, 2016).
Moreover, almond growing has also shifted to subtropical regions,
such as Central Morocco (Kodad and Socias i Company, 2014) or
southern Peru (J. Torrents, unpublished), where the question of
adaptability of the recently released late-blooming cultivars has arisen
due to the possible lack of sufficient chill during winter dormancy. The
traditional almond cultivars are considered to be well adapted to the
conditions of mild winters with low chilling accumulation, such as
California (Brown, 1952) and Tunisia (Crossa-Raynaud, 1955), but
some of the new cultivar releases show very high chilling requirements
as compared to the traditional cultivars (Socias i Company et al., 2015).
Consequently, almond growing is at present expanding towards two
opposite environmental conditions: cold regions with high risk of frosts
and warm regions with very low chilling accumulation. Additionally,
the new environmental conditions arisen by the actual climatic changes

⁎
Corresponding author.
E-mail address: jmalonsos@aragon.es (J.M. Alonso Segura).

http://dx.doi.org/10.1016/j.scienta.2017.05.036
Received 11 April 2017; Received in revised form 29 May 2017; Accepted 30 May 2017
Available online 07 June 2017
0304-4238/ © 2017 Elsevier B.V. All rights reserved.
J.M. Alonso Segura et al.
Fig. 1. Spanish almond kernel production in the period 2001–2016 (Cooperativas Agroalimentarias).
must be taken into account when considering the new scenario of al-
mond growing (Socias i Company et al., 2003). Thus, almond breeders
have to consider the adaptability of almond to these different condi-
tions depending on the chilling and heat requirements of the cultivars
and to point out the need of establishing clear objectives for blooming
time in the almond breeding programmes.
2. Almond blooming
Almond was traditionally the first flowering species among all cul-
tivated Prunus. However, since late blooming is a desirable trait in
breeding programmes, significant breeding progress has been made in
the delay of the bloom time with the newly released cultivars. As a
consequence, almond cultivars probably show the widest range of
blooming dates among all the fruit and nut species (Socias i Company
and Felipe, 1992). Blooming takes place when the petals appear
showing the different flower elements (Felipe, 1977a). In the almond
germoplasm collection of Zaragoza (Spain) bloom starts in the middle
of January for cultivars of North African origin, with very low chilling
requirements, and continues until the end of March or early April for
the extra-late blooming releases and breeding selections.
Bloom date is considered dependent on the progression of winter
temperatures affecting the different stages of dormancy (Lang et al.,
1987). At dormancy onset, flower organogenesis is complete (Lamp
et al., 2001). During endodormancy, bud growth is constant but barely
perceptible. Low temperatures are the most important factor affecting
endodormancy completion, though other climatic factors such as heat,
photoperiod, light intensity, and mist humidity can also affect en-
dodormancy development (Freeman and Martin, 1981). After accu-
mulating sufficient chilling, endodormancy concludes, giving way to
ecodormancy. The end of endodormancy is considered to coincide with
the completion of the chilling requirements and cannot be sustained by
any visible change. During ecodormancy, flower bud growth mostly
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Scientia Horticulturae 224 (2017) 61–67
depends on heat accumulation. Accumulation of heat units is important
in the active growth processes of most plants, including breaking of
endodormancy (Sherman and Rodríguez-Alcazar, 1996). Once ecodo-
mancy has started, bud growth rate depends on favourable growth
temperatures and can increase considerably with warm temperatures
during the final stages of winter in the Mediterranean climates (Kester
and Gradziel, 1996; Westwood, 1978). The ecodormancy is considered
complete at F50, or anthesis of the 50% of the flowers (Tabuenca et al.,
1972), a stage easily estimated in the field through the record of the
phenological stages of buds and flowers (Felipe, 1977a). The F50 stage is
thus considered to be the stage of completion of the heat requirements
for bud development. Most almond cultivars show rather low chilling
requirements, although high variability is observed among cultivars
(Tabuenca, 1972; Alonso et al., 2005). Similarly the almond heat re-
quirements are quite low and very variable (Tabuenca et al., 1972;
Alonso et al., 2005).
Blooming date may change from year to year, depending on the
climatic conditions of the year (Tabuenca and Herrero, 1966; Rattigan
and Hill, 1986) conducing to break endo- and ecodormancy (Lang et al.,
1987) after fulfilling the chilling and the heat requirements of the
flower buds.
Although the blooming sequence of the different cultivars is rela-
tively constant over the years, small variations in the order of blooming
may occur depending on the location and the temperatures of each
season (Felipe, 1977b; Dicenta et al., 1993; DeGrandi-Hoffman et al.,
1996), since the evolution of temperatures during winter dormancy
may lead to different dosing regimens of chilling and heat accumulation
during the whole dormancy period of every year resulting in these
variations (Alonso et al., 2005).
Similarly, the duration of bloom is dependent on each cultivar and
on the temperature during the blooming season. Thus, the flower
opening sequence of some cultivars is very short, as in ‘Ferraduel’,
lasting a few days, independently of the temperature, whereas in others
J.M. Alonso Segura et al.
is very long, as in ‘Bertina’, being always longer at low temperatures
and shorter at higher temperatures and following closely a normal
distribution (Socias i Company, 1983; Hill et al., 1985; Socias i
Company and Felipe, 1987; Bernad and Socias i Company, 1995). In
any case, the progression of almond bloom is determined by the accu-
mulation of heat units (DeGrandi-Hoffman et al., 1996).
3. Chilling and heat requirements
Only a few reports have examined chill and heat requirements in
almond, probably because almond has low chill requirements and
currently this is not a limitation for this crop. Brown (1952) and Crossa-
Raynaud (1955) reported that almond had lower chilling requirements
than the other Prunus species since it was well adapted to warm winters.
Most almond cultivars show rather low chilling requirements, although
high variability is observed among cultivars (Tabuenca, 1972; Alonso
et al., 2005). Similarly, the almond heat requirements are quite low
though very variable (Tabuenca et al., 1972; Alonso et al., 2005). The
first estimates of chilling requirements for dormancy break in almond
were quantified by Tabuenca (1972) in 15 cultivars and local Spanish
clones as the accumulation of hours under 7 °C (44.6 °F) and 12 °C
(53.6 °F) from dormancy onset until endodormancy break, a critical
point determined by the observation of a significant increase of the dry
weight of the buds (Brown and Abi-Fadel, 1953). The chilling re-
quirements thus obtained ranged from 90 to 427 h bellow 7 °C and from
422 to 940 h bellow 12 °C. In another work of the same year (Tabuenca
et al., 1972), the first estimations of the heat requirements for blooming
were stabilised in 38 almond cultivars, being quantified as the accu-
mulation of maximum and minimum daily temperatures from dor-
mancy break until full bloom. In this case, the endodormancy break
dates were estimated through a statistic method based on correlations
between temperatures and bloom dates using data accumulated over
several years. The cultivars with lower heat requirements showed totals
ranging between 470–572 °C for the maximum daily temperatures and
152–202 °C for the minimum daily temperatures. Cultivars with higher
heat requirements showed totals of 1104–1331 °C, and 373–513 °C re-
spectively.
Several years after the first studies of Tabuenca in almond, re-
searchers from the University of Utah developed a method to quantify
the chill unit accumulation coinciding with the completion of en-
dodormancy, as well as the growing-degree-hour (GDH) accumulation
to full bloom (Richardson et al., 1974,1975). This method has become
over the years the mostly adopted for measuring the chilling and heat
requirements of fruit tree species. The chill unit model (Richardson
et al., 1974) quantifies the chilling requirements as well as the relative
effectiveness of various temperatures for releasing dormancy, and is
supported by controlled environment studies on the influence of par-
ticular temperature ranges on dormancy (Erez and Lavee, 1971). The
phenological model for flower bud development of fruit trees
(Richardson et al., 1975) consist in a linear model with a base tem-
perature of 4.5 °C and an upper limit of 25 °C, thus the greatest heat
accumulation for any 1 h is 20.5 GDH’s. The use of computers fa-
cilitated the development of more accurate quantification of chill and
heat requirements, as well as the assessment of the theoretical en-
dodormancy end date using regression and other statistical techniques.
Rattigan and Hill (1986) applied the systematic trial-and-error statis-
tical approach of Ashcroft et al. (1977) and evaluated the chilling and
heat requirements of 12 almond cultivars grown in the mild winter
climate of Australia where they also observed a significant location
influence on the blooming behaviour of three cultivars (Rattigan and
Hill, 1987). These researchers found early completion of chill require-
ments in almond and low variability between cultivars (220–350 Utah
Chill Units) and a larger variability for heat requirements (5300–9000
GDH), suggesting a major effect of the heat requirements on de-
termining the final bloom date.
More recently, Egea et al. (2003) estimated the chilling and heat
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Scientia Horticulturae 224 (2017) 61–67
requirements of 10 almond cultivars and breeding selections grown in
the mild winter climate of southeastern Spain by assessing the en-
dodormacy breaking date using the traditional bud weight method. In
this study, almond chilling requirements for flowering showed a very
high variability, ranging from 266 to 996 CU, whereas the heat re-
quirements showed less variability, ranging from 5942 to 7438 GDH.
A more recent quantification of chill and heat requirements for
flowering in almond has been reported by Alonso et al. (2005). This
work can be considered an extension of the work done by Tabuenca
et al. (1972), which was carried out in the same geographical location,
Zaragoza (Spain), but improving the regression method for determi-
nation of the end of endodormancy. Alonso et al. (2005) found that in
the cold winter climatic conditions of the Ebro Central Valley of Spain,
almond showed chilling requirements ranging between 350 and 500
CU, and heat requirements ranging between 5500 and 9300 GDH.
The avoidance of cold injury is often accomplished by increasing
either chilling requirement or post-chill heat requirement in order to
delay bloom. It is evident that the Spanish cultivars ‘Felisia’, ‘Moncayo’,
‘Guara’, ‘Cambra’, ‘Aylés’ and ‘Masbovera’, and the French cultivars
‘Ferragnès’, ‘Ferralise’ and ‘Lauranne’, obtained in breeding pro-
grammes, show high heat requirements. On the other hand, traditional
cultivars such as ‘Desmayo Largueta’, ‘Marcona’, ‘Ramillete’,
‘Constantini’ and ‘Pou de Felanitx’ show low heat requirements.
If a cultivar gets too much chilling, either because it has been
planted in a colder climate than it is adapted to, or because of unusually
cold winters in a traditional growing region, it may be subject to early
bloom with possible damage by later spring frosts. If a cultivar gets too
little chilling, flowering and fruiting are delayed, the tree may not have
sufficient fruit set for a commercial crop, while leafing is delayed and
erratic.
Since almond chill requirements are quite low, heat requirements
are a greater determinant than chilling requirements in regulating the
blooming date in climates characterised by cold temperatures during
dormancy. Flower buds are thus exposed to an excess of chilling, with
continuous low temperatures with high chilling effectiveness.
Consequently, the endodormancy phase is shortened and the chilling
requirement is accomplished in a very short period of days in most
almond cultivars (Rattigan and Hill, 1986; Alonso et al., 2005). On the
other hand, completion of heat requirements is very slow due to the
very scarce occurrence of favourable temperatures for vegetative
growth, resulting in a low effectiveness in the completion of heat re-
quirements during ecodormancy. Tabuenca et al. (1972) explained that
almond genotypes growing in climates where they are saturated with
chill units during winter start the accumulation of heat units at the
same time, beginning with spring warming and, as a consequence, the
final bloom date is strongly correlated with the heat requirements of
each genotype. Consequently, a shortening of the blooming season to
only four weeks is observed between the first and the last blooming
cultivars. However, the pattern of completion of chill and heat re-
quirements in climates characterised by mild temperatures during
dormancy is just the opposite. The quality of temperatures to complete
chill is very limiting, whereas the quality of warm temperatures to
promote growth is very high, resulting in chilling requirements being
the major determinant of bloom dates in these conditions (Egea et al.,
2003).
The chilling requirements of any cultivar are not related to its
blooming time as seen in Table 1, where several cultivars are ordained
according to their blooming date in the Spanish almond reference
collection of the CITA of Aragón in Zaragoza. Thus, the cultivar with
the lowest chilling requirement, ‘Felisia’, is a late-blooming cultivar, as
also happens with ‘Tardive de la Verdière’.
While the succession of bloom timing among the different almond
cultivars is generally maintained over the years (Felipe, 1977b), small
shifts may result from subtle cultivar differences in endodormancy and
ecodormancy interactions (Couvillon and Erez, 1985; Gilreath and
Buchanan, 1981). As a consequence, unless two cultivars have identical
J.M. Alonso Segura et al. Scientia Horticulturae 224 (2017) 61–67

Table 1 4. Genetics of blooming time

Chilling and heat requirements of several almond cultivars with different blooming times
(Alonso et al., 2005; Alonso and Socias i Company, 2009; Socias i Company et al., 2015).
With the purpose of reducing the problem of spring frosts, late
Cultivar Country of Average Chilling Heat flowering has been an objective of most almond breeding programs in
cultivar blooming requirements requirements order to have flowering when the risk of frosts is low or non-existent
origin date in (CU)z (GDH)y (Socias i Company et al., 2012). Moreover, later in the season, tem-
Zaragoza peratures are usually higher and more favourable for the processes of
Zahaf Tunisia 9 Feb 392 5611 pollination and fertilization (Kester and Asay, 1975). The large varia-
Desmayo Spain 11 Feb 428 5458 bility of blooming date of the different almond cultivars has provided
Largueta almond breeders a wide possibility of selecting parents for the breeding
Marcona Spain 20 Feb 428 6603 crosses. This trait is considered to be quantitatively inherited in most
Nonpareil USA 27 Feb 403 7758
fruit species (Anderson and Seeley, 1993) and most results on the
Belona Spain 1 Mar 353 7741
Soleta Spain 1 Mar 340 7872 transmission of flowering time in almond show this quantitative be-
Cristomorto Italy 3 Mar 428 8027 haviour (Kester, 1965; Grasselly and Gall, 1967; Grasselly, 1972;
Aï France 4 Mar 444 8051 Vargas and Romero, 1988) with a range of heritability estimates from
Ferragnès France 4 Mar 444 8051
0.804 (Kester et al., 1977) to 0.67 (Dicenta et al., 1993), a difference
Guara Spain 4 Mar 340 8159
Tuono Italy 4 Mar 463 7978 that probably results from the wider genetic base of the parents in-
Tardive de la France 5 Mar 358 8814 volved in the later study. In extreme cases, especially when both par-
Verdière ents are late flowering, only a few genotypes of the offspring would be
Primorskij Ukraine 6 Mar 428 8434 expected to be later blooming than both parents, thus increasing the
Titan USA 7 Mar 444 8457
difficulty of retarding bloom in each generation (Grasselly and Gall,
Felisia Spain 12 Mar 329 9465
Vialfas Spain 21 Mar 503 10066 1967).
Mardía Spain 24 Mar 503 10233 Despite these quantitative estimates, Kester (1965) suggested that in
some progenies of the late-blooming budsport ‘Tardy Nonpareil’, a
z
Chilling units. single dominant gene could be involved in determining the date of
y
Growing Degree Hours in °Celsius.
flowering, since a bimodal distribution of blooming classes was ob-
served among the seedlings obtained. Similar results were also obtained
heat and chilling requirements, their bloom overlap will depend mainly
with the same cultivar by Grasselly (1978), although this work was
on the temperature accumulation during dormancy. With traditional
partially discontinued because of the usual correlation of late bloom
commercial cultivars, most of them self-incompatible, the association of
and low bud density observed in these progenies (Grasselly and Olivier,
different inter-compatible cultivars with synchronised bloom time is
1985). However, some of the selections obtained in this French
essential to ensure inter-pollination, and thus the obtaining of an ac-
breeding programme have been included as parents in other pro-
ceptable commercial yield. In some subtropical regions, the blooming
grammes from which some selections have been released (Dicenta
date of these cultivars is asynchronous when the winter temperatures
et al., 2009).
are low, when the differences in heat requirements greatly affect
The same bimodal distribution was observed in the Zaragoza
blooming time, resulting in low crops (Kodad and Socias i Company,
breeding programme with another parent than ‘Tardy Nonpareil’, al-
2014). The incidence of low temperatures following the start of bloom,
though from its descent. The first cross included ‘Titan’, an open pol-
will result in a prolonged bloom, whereas warmer than normal tem-
linated seedling from ‘Tardy Nonpareil’ selected because of its late
peratures results in the compression of the bloom season (Chandler and
flowering date in order to produce natural hybrids when being cross-
Brown, 1951), since flower development is dependent on the tem-
pollinated by the peach ‘Nemaguard’ in isolated mixed orchards, since
perature (Rattigan and Hill, 1987; Bernad and Socias i Company, 1995).
self-pollination of ‘Tardy Nonpareil’ would not produce any fruit
Another cause of bloom variation is the base temperature required
(Jones, 1972). From the progeny ‘Titan’ × ‘Tuono’ an individual was
for flower bud development, which differs not only among species
released as ‘Felisia’ (Socias i Company and Felipe, 1999), the latest
(Richardson et al., 1975; Rattigan and Hill, 1987), but also among
blooming cultivar at that time, which was used as parent in several
cultivars within a species, including almond (Bernad and Socias i
crosses where a bimodal distribution was observed (Socias i Company
Company, 1995; DeGrandi-Hoffman et al., 1996; Hill et al., 1985).
et al., 1999). These progenies allowed the identification of a major gene
There is genetic variability for the optimum temperature at which
named Lb (late bloom), which delayed bloom 15 days on the average
chilling takes place. It has been suggested that low chill cultivars have a
(Socias i Company et al., 1999). This allele was soon mapped in the
higher optimum temperature for chill unit accumulation than high chill
linkage group 4 (Ballester et al., 2001), which can be traced not only
cultivars (Gurdian and Biggs, 1964). In addition, each cultivar could
back to a single mutant, ‘Tardy Nonpareil’, but also further to the fol-
react in a different way to occasional extreme temperatures or to a
lowing generations (Socias i Company et al., 2006).
sudden alternation of cold and warm temperatures, a typical phenom-
The presence of this allele shows that the time of almond flowering
enon occurring during the spring in temperate climates (Couvillon and
in these progenies is both a quantitative and a qualitative trait, with late
Erez, 1985; Erez and Couvillon, 1987).
bloom dominant over early bloom (Socias i Company et al., 1999). The
Prunus species develop vegetative and flower buds separately with
quantitative analysis confirmed that this allele explained at least 79% of
flowering usually taking place before leafing. Thus, it has been specu-
the phenotypic variation (Ballester et al., 2001), since the qualitative
lated that flower buds have lower chilling requirements than leaf buds,
effect of this allele was quantitatively modified by minor genes.
or that leaf buds have higher heat requirements than flower buds. In
Grasselly and Olivier (1985) described the possible linkage between
contrast, at given locations, some genotypes with low chill require-
late bloom and low productivity in crosses involving ‘Tardy Nonpareil’.
ments leaf out simultaneously with or just before flowering. It may be
This linkage could be due to the Californian origin of ‘Tardy Nonpareil’,
that leaf buds start accumulating heat units at a lower threshold than
since Californian cultivars, such as ‘Nonpareil’, have shown lower
flower buds; consequently, during prolonged cool weather leaf buds get
flower bud density than traditional Spanish cultivars such as ‘Marcona’,
ahead of flower buds in heat accumulation and thus begin breaking
and new releases such as ‘Guara’ (Socias i Company, 1988). A high
dormancy earlier.
flower bud density has been shown to increase the possibilities to en-
dure late frosts during bloom (Kodad and Socias i Company, 2005)
because the presence of a high number of flowers increases the

64
J.M. Alonso Segura et al.
possibilities of survival of some flowers after a frost and, thus, allowing
an acceptable crop. Frost risks are lower in California than in Spain,
thus leading to the prevalence of cultivars with low or medium flower
bud density, probably to reduce the problems related to flower quality
(Socias i Company and Felipe, 1994), whereas in Spain, where frost
risks are important (Felipe, 1988), a high flower bud density has been a
positive trait for cultivar evaluation in order to ensure a crop (Kodad
and Socias i Company, 2008). The same hypothesis has been put for-
ward to explain the high flower bud density of some peach cultivars
from the United States and Canada (Okie and Werner, 1996; Werner
et al., 1988), and Hungary (Szabó et al., 1998).
5. Almond blooming and breeding
Late bloom has been an objective of almond breeding, although
with much more attention in the Mediterranean region than in
California (Grasselly, 1972; Rikhter, 1972; Kester and Asay, 1975;
Socias i Company et al., 2012). In the years 1960s, cultivars then
considered as late blooming spread in most Mediterranean growing
regions, including both traditional and improved cultivars. The tradi-
tional cultivars were mostly from Puglia, such as ‘Tuono’ and ‘Cristo-
morto’ and, to a lower proportion the French ‘Aï’ and ‘Tardive de la
Verdière’. The improved cultivars were mostly from the French
breeding programme, such as ‘Ferragnès’ and ‘Ferraduel’ and, to a lesser
extent, the then Soviet ‘Miagkoskorlupyj’, ‘Primorskij’ and ‘Yalstinskij’.
Both ‘Tuono’ and ‘Ferragnès’, of similar blooming time, reached and
still maintain a relevant presence in many Mediterranean orchards.
When the date of full flowering was established by the almond de-
scriptors (Gülcan, 1985), the scale ranged from 1 for the very early
blooming ‘Cavaliera’ to 9 for ‘Tardy Nonpareil’, then considered as
extremely late. In this scale, ‘Ferragnès’ is rated as very late, with the 8
score. Considering the blooming time of some of the new releases
(Socias i Company and Felipe, 1999; Socias i Company et al., 2008a,
2015; Vargas et al., 2008; Dicenta et al., 2009), later or much later than
‘Tardy Nonpareil’, this present scale has become inoperable. Conse-
quently, ‘Ferragnès’ could be considered as a medium-blooming cul-
tivar, and at the GREMPA meeting of Meknès, Morocco, in 1997 it was
suggested to reduce this scale and lower ‘Tardy Nonpareil’ to the rating
of 6, leaving the higher ratings of 7, 8 and 9 for later blooming culti-
vars.
The first attempts to introduce late bloom as an objective in almond
breeding was that of A.A. Rikhter at the Nikita Botanical Garden of
Yalta, Crimea, then the Soviet Union. The harsh conditions of the al-
mond growing regions in Central Asia, Southern Russia and Ukraine
where almond was intended to be grown, made late-bloom and frost
resistance the main requirements of this breeding programme (Rikhter,
1972). Late-bloom was linked to stable dormancy in order to avoid
bloom after short periods of warm temperatures, called by Rikhter
“provocative thaws”. Thus, an indirect selection for high heat require-
ments was carried on, because the deep rest during eco-dormancy (Lang
et al., 1987) lasted for a long time, and only after several periods of
“provocative thaws”, when frosts would not be frequent, the plants
would bloom. This trait was selected in the southern seashore of Crimea
where these “provocative thaws” are frequent, being considered a ty-
pical region for this kind of temperature evolution. Consequently, the
best known releases from this breeding program, ‘Primorskij’, ‘Miag-
koskorlupyj’ and ‘Yaltinskij’, were characterised by low chilling re-
quirements but very high heat requirements (Tabuenca et al., 1972;
Alonso et al., 2005). Additionally, the best selections showing stable
late-bloom were tested for frost resistance in the Simferopol region
where winter temperatures may be as low as −20 °C (–4 °F). These
varieties were tested in the then Soviet Union with varying success
(Rikhter, 1972) and also introduced in southern Europe, where they
have not attained real importance because of their growth habit, but
have been introduced as parents in several breeding programmes
(Vargas and Romero, 1988; Dicenta et al., 2009).
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Scientia Horticulturae 224 (2017) 61–67
The following attempt was carried out by C. Grasselly in France,
firstly taking ‘Tardy Nonpareil’ as a source for late bloom (Grasselly,
1978). However, as mentioned above, this approach was partially dis-
continued because of the usual correlation of late bloom and low bud
density, although this linkage could have been broken in some cases
(Grasselly and Olivier, 1984). Consequently, the late blooming releases
of the French breeding program have been obtained from crosses in-
volving the late blooming French cultivar ‘Aï’ and cultivars from the
Italian region of Puglia, a region characterised by the presence of late-
blooming cultivars, including ‘Ferralise’ (Grasselly and Crossa-
Raynaud, 1980) and ‘Mandaline’ (Duval, 1999).
The CITA breeding program firstly looked for late bloom in crosses
involving French and Puglian cultivars, such as in ‘Moncayo’ (Felipe
and Socias i Company, 1987), originated in the cross ‘Tuono’ × ‘-
Tardive de la Verdière’. However, this program later introduced ‘Titan’,
a seedling of ‘Tardy Nonpareil’, where the linkage between late bloom
and low productivity had also been apparently broken. Thus, the first
late-blooming release, ‘Felisia’ (Socias i Company and Felipe, 1999) was
obtained in the cross ‘Titan’ × ‘Tuono’ and the latest releases, ‘Mardía’
(Socias i Company et al., 2008a) and ‘Vialfas (Socias i Company et al.,
2015) from “Felisia” × ‘Bertina’, a late-blooming local selection of
unknown origin (Felipe, 2000). These last cultivars are characterised by
a permanent late-blooming as determined by very high requirements of
both chilling and heating (Table 1). At present, most crosses involve a
parent derived from ‘Tardy Nonpareil’, thus possessing the major allele
Lb (Socias i Company et al., 2006).
The IRTA and the CEBAS breeding programmes have included
mostly the Crimean release ‘Primosrkij’ in their crosses with the ob-
jective of late blooming, as well as Italian cultivars from Puglia and
French releases, both named cultivars and unnamed selections such as
R1000 which is a seedling of ‘Tardy Nonpareil’.
No breeding programme has included early blooming as on objec-
tive since most traditional almond cultivars, as already commented,
showed low chilling requirements (Table 1) and only the new releases
with very high chilling requirements may show problems of lack of
chilling in subtropical regions or in new scenarios arisen by the present
climatic changes. Early-blooming cultivars, however, really need to be
incorporated in almond breeding programmes because they mostly lack
two of the most important traits defined as objectives in the present
programmes (Socias i Company et al., 2012): self-compatibility (Socias i
Company, 1990) and kernel quality (Socias i Company et al., 2008b).
Early blooming was considered an interesting trait for almond
growing in arid regions, where early blooming could involve also early
ripening, before the extreme summer climatic conditions of heat and
drought. Since many wild almond species show a very short fruiting
period, the utilisation of these wild species was suggested in almond
breeding programmes (Grasselly, 1976), although their utilisation has
been limited and not for this objective (Gradziel, 2003).
6. Conclusion
The objective of retarding blooming time in almond has been at-
tained, as shown by the release of very late blooming cultivars from
different breeding programs (Table 2). These advances have shown that
the use of late-blooming cultivars with distant geographical origins
(Table 3) has allowed the accumulation of minor genes with additive
effects. All these releases have received an important contribution from
the Puglia genetic pool, with a minimum of 25% in ‘Mardía’ and ‘Tar-
dona’, but the contribution of some local selections from Spain, iden-
tified by their late bloom, such as ‘Bertina’, has been also important. In
some releases, including ‘Felisia’, ‘Mardía’, ‘Vialfas’ and ‘Tardona’, this
accumulation has been stressed by the effect of the major allele Lb.
Taking into account that the Californian gene pool is mostly of French
origin (Kester et al., 1990) and that most parents in the Yalta breeding
program were from France and the USA (Rikhter, 1972), the con-
tribution of the French genepool to these releases is also important. This
J.M. Alonso Segura et al. Scientia Horticulturae 224 (2017) 61–67

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