e Scientia Letters ESL/ 2021/Feb/ Vol9
Introduction Therefore depleted local resource
storage following heavy flowering year,
Irregular or alternate bearing is a major
only leaves are produced. Leaves have
problem for sustained productivity of
lower resource demands than fruit and
many fruit crops, particularly in
participate in increased local storage for
changing climate scenario. It refers to
the following year when flowers/fruits
the propensity of an entire tree to
are produced.
produce more than the average crop in
one year, followed by less than the
Furthermore, growing fruits are known
average crop in next year. It is a big
to suppress developing buds that
economic issue for the growers, as the
would otherwise transition to flower
net return to the farmer during the off-
buds in the next year. As a result,
year crop is low, and it disrupts the
during years with greater than average
orchard maintenance cycles. Long-
fruit production, the majority of
term studies of the development of
developing buds are suppressed so
flowers and fruits have shown that
that the following year only a few
individual shoots alternate between
flowers and fruit are produced. This
years when flowers and fruits are
results in regular oscillations between
produced and years when only leaves
high and low yields in alternating years.
are produced. (Goldschmidt, 2013;
(Hoblyn et al., 1936) gave an index that
Sharma et al., 2015). The theory that
measured the severity of variance
alternate bearing in individual tree
during consecutive years in yield in fruit
occurs as a result of variations in local
crops. Then this index was called as the
resource storage over years is
Biennial/Alternate Bearing Index by
supported by two observations:
Monselise and Goldschmidt, 1982 and
1. Individual outgrowth, not
they gave following formula for AB
branches or whole trees, gave
quantification:
rise to alternate leaf and fruit
Formula 1: Alternate bearing index (I) =
production each year.
(year 1 yield) - (year 2 yield) / (year 1
2. The majority of sugars produced
yield + year 2 yield), where I = 0 is no
by a leaf are translocated for
alternate bearing and I = 1.0 is
only short distances to the
complete alternate bearing;
adjacent fruit or branch tissues.
15
e Scientia Letters ESL/ 2021/Feb/ Vol9
Formula 2: Alternate bearing (AB) = (Smith et al., 2004), pecan (Wood et al.,
(current year's yield) - (5-year running 2004), and pistachio (Rosenstock et al.,
average yield)/5-year running average 2010). Later modified to “Modified
yield; when the current yield is 20% less Alternate Bearing Index” (MABI), to
than the 5-year average, it is an off-crop. assign role of flowering in the
The Biennial Bearing Index has been phenomenon. An image of vegetative and
used in various fruit crops like in apple floral bud is shown in Fig.1, which is under
(Barritt et al., 1997), mango (Reddy et al., inductive control of weather parameters.
2003), coffee (Cilas et al., 2011), citrus
Fig.1 Mango: (A) Vegetative and (B) Floral bud
Impact of intrinsic and exogenous In general, tropical trees are induced to
factors on bearing habit flower by environmental signals.
Vegetative and flower bud formation in Besides these there are several
tropics and subtropics varies for same endogenous and exogenous factors
fruit species, and implicates age of responsible for this rhythm.
shoot and cool inductive temperatures Endogenous factors can be classified
programmed by the coincidence of into: Genotypic differences, effects of
internal and environmental signals in leaf number and maturity, competition
higher plants. Four major pathways between vegetative and reproductive
have been postulated for induction of sinks, crop overload, carbohydrate
flowering: environmental induction reserves (C:N), tree age and vigour,
through i) photoperiod and ii) hormonal imbalance (auxin, GAs, CK,
temperature, iii) autonomous floral ABA and polyamines), natural
initiation( regulated by age and maturity abscission of buds, flowers and fruits,
of shoot) , and iv) regulation by flower inhibition by growing fruits,
gibberellins (or anti-gibberellins). nutrient status, pollination, age and size
of shoot, fruit drop, fruiting habit, root
16
e Scientia Letters ESL/ 2021/Feb/ Vol9
stocks, growth pattern, branch cause of the phenomenon of biennial
anatomy and others. Exogenous bearing. It is quite probable that an
factors consists of environmental accumulation of these compounds may
factors, temperature, light, relative create favourable conditions for the
humidity, rains and snow (temperate synthesis and action of the substances
species), spring frost, hails, water responsible for flowering.
stress, edaphic factors, biotic factors 3. Hormonal Balance: Flowering time is
etc. few are discussed for a largely affected by large chemical
comprehensive understanding: constituents such as plant hormones
1. Growth Pattern: Growth in fruit trees (Davis, 2009; Domagalska et al., 2010).
usually takes place in different flushes, Biennial bearing problem appears to be
which vary in different parts of the closely associated with the fruit
geography. Early initiation and growth development process and the inhibitory
cessation, followed by a definite influence of the developing fruits on
dormant period, will help the shoots to vegetative growth. Higher levels of
achieve the proper physiological auxin like substances and an inhibitor
maturity that is necessary for the (similar to ABA) and lower levels of
initiation of the fruit-bud. The growth of Gibberellin (GA3) like substances are
shoots, however, is solely a varietal vital for a florigenous shoot. There is
feature and is regulated by the on and inverse relationship between the level
off-year process of the trees in biennial of endogenous inhibitor in the shoot
bearing varieties, rather than by the and vegetative growth. Higher inhibitor
time of origin and cessation of shoot content promotes flowering.
growth. In an On year, flower buds are 4. Environmental Triggers: Weather
marked by shoots of any size or conditions impact flower and fruit
maturity, whereas in a year Off, even development and can exacerbate the
the shoots of the appropriate size and alternate bearing tendency of individual
maturity fail to flower. tress or whole orchards. For example, a
2. Carbon / Nitrogen Ratio: In fruit severe frost during flowering would kill
plants, N and CHO reserves play developing flower tissue and result in a
important role in flower bud initiation, very low fruit set as is happening in
even if these do not form the primary mango in subtropics. As a result of the
17
e Scientia Letters ESL/ 2021/Feb/ Vol9
low fruit set, and reduced use of stored orchard, pattern of alternate bearing.
carbohydrates at each individual shoot, Adverse climatic conditions like rain,
the following year most shoots will high humidity and low temperature
produce flowers and set fruit. If all sometimes convert an on year into off
outgrowths on a tree are synchronized year directly or by promoting the
to flower during the same year it will incidence of diseases like powdery
generate a whole tree, and even whole mildew, anthracnose and poor fruit set.
Table No. 1: Weather attributes associated with flowering in perennial fruit crops
Crop Critical weather parameters for floral induction References
Mango Temperature below 15–20 °C, with florally inductive Nunez-Elisea and
(subtropical temperatures varying between cultivars. Cool Davenport, 1994,
& tropical) temperature is the dominant induction factor ,under Whiley et al., 1989,
subtropical conditions. Under tropical conditions, in Pandey, 1989
which cold inductive temperatures may be brief,
erratic or non-existent in some season.
Avocado Period of low temp around 150 /100, 180 /15 0 C Wakabayashia and
Huber, 2001
Litchi Temp around 100C, low temperature induces Menzel, 1983
flowering
Apple Most of apple varieties require 1000-1600 chilling Wilkie et al., 2008
hours during winter to break the rest period, however
some low chill varieties require only 500-800 chilling
hours.The average summer temperature should be
around 21-240C during active growth period, Shading
30% to 70%
Peach Chilling requirement is necessary for flower induction Scarza and Sherman,
1996
Orange Flowering occurs in the spring after several months Hart et al., 1991
with mean temp. below 15°C
18
e Scientia Letters ESL/ 2021/Feb/ Vol9
Synchronization of alternation over shoots for the next season and their
wide areas often has been blamed on subsequent fruit-bud differentiation.
spring frost with apples, olives Therefore, the fruit load appears to be
(Morettini, 1950), pecans (Sparks, one of the main conditioning factors for
1974), and mangos in cool areas (Singh ‘on’ or ‘off’ year in fruit crops. This
et al., 1974). There are two major hypothesises, that if fruit load is more,
differences which exist between then it blocks the recognition of
tropical and temperate deciduous flowering inductive signal (s). It
horticultural trees when it comes to prevents the emergence of
floral initiation. Tropical tree species inflorescence and bud break (Albrigo
such as mango initiate flowering in and Saúco, 2004; Verreynne and
response to an environmental stimulus, Lovatt, 2009).
while the temperate deciduous species,
such as apple, initiate flowers Cultural Practices to overcome
autonomously. Secondly, temperate Alternate Bearing
deciduous horticultural trees undergo a 1. Use of chemicals like paclobutrazol:
period of dormancy between floral Hormonal regulation of flowering is
initiation and anthesis, while in tropical one of key mechanisms responsible
species, including mango, floral for induction of flowering in fruit
development is continuous from floral trees, which is based on a balanced
induction to anthesis (however ratio of hormonal synthesis and
difference exists for subtropics) (Table transport to the terminal shoots.
1). Paclobutrazol (PBZ) treatment is a
5. Crop Load: Generally, moderate technological intervention that
blossoming is one of the chief suppresses gibberellins and
conditions of annual fruit bearing in fruit promotes flowering in mango under
trees. Fruiting is an exhausting process influence of various
and the number of fruits retained till hormones Paclobutrazol checks.
harvest is a varietal feature. The total
number of fruits that are harvested is
important because of their deleterious
influence on the production of new
19
e Scientia Letters ESL/ 2021/Feb/ Vol9
A) Control B) Treated
Fig.2 Effect of PBZ on flowering.
2. Gibberellic Acid biosynthesis and 5. Early harvesting: It was shown that
increased cytokinin level, chlorophyll removal of fruits from ‘On’ year crop
content improves mineral uptake and enhanced the return bloom (Shalom
carbohydrate gradient of the entire et al., 2014). Similar techniques have
plant system. This helps in achieving a been used with mango and apples
balance in C: N ratio which induces (Moss et al., 1977).
flowering normally. The field should be 6. Use of rootstocks- Response of
irrigated immediately after treatment to mango varieties at different height of
increase efficiency. Effect of PBZ grafting on rootstock was studied by
application to enhance and regulate (Pandey, 1989). Similarly, effects of
flowering in mango field is well clonal rootstocks on Hass avocado
documented. (Fig.3). on alternate bearing has been
3. Deblossoming: Deblossoming of the elucidated by (Mickelbart et al.,
panicles with NAA @ 200ppm during 2007).
‘on’ year. NAA has been used with 7. Planting of fairly regular bearing
success on mandarin-type fruits varieties like in mango: Amrapali,
(Galliani et al., 1975). Banganpalli, Bangalora and Neelum
4. Girdling: The process of girdling in are suggested for getting regular
citrus in autumn increases the fruits.
flowering next spring, while when it is 8. Pruning: Removing bearing surface
done at blossom time, it increases the (fruit buds) stimulates vegetative
set. Similar practices are also used growth from remaining buds. This
with ‘Nabal’ avocado, where inducing promotion of vegetative growth
alternation on half of the tree proves inhibits many of the remaining buds
commercially beneficial. from becoming floral.
20
