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Bee World
ISSN: 0005-772X (Print) 2376-7618 (Online) Journal homepage: http://www.tandfonline.com/loi/tbee20
The Basic Concept of Honey Bee Breeding
Programs
A. Uzunov, E. W. Brascamp & R. Büchler
To cite this article: A. Uzunov, E. W. Brascamp & R. Büchler (2017) The Basic Concept of Honey
Bee Breeding Programs, Bee World, 94:3, 84-87, DOI: 10.1080/0005772X.2017.1345427
To link to this article: http://dx.doi.org/10.1080/0005772X.2017.1345427
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ARTICLE
Page 84 • VOL 94 • September 2017 • Bee World
The Basic Concept of Honey Bee
Breeding Programs
A. Uzunov, E. W. Brascamp and R. Büchler
Selective honey bee breeding is a phe-
nomenon that fascinates beekeepers
around the world. ey oen regard it as
one of the most enigmatic and complex
aspects of beekeeping. Indeed, according
to our experiences participating in many
international projects, both beekeepers
and bee experts without a background
in plant or animal breeding oen have
trouble correctly interpreting and con-
ceptually visualizing the breeding process.
ese diculties arise partly because
of the complex reproductive biology of
honey bees, where queens mate with a
multitude of drones. Fundamentally the
greatest struggle for people to understand
is how selection of animals with preferred
characteristics in one generation leads to
improved progeny in the next.
e leading misconception regarding
honey bee breeding is confusing breeding
with the simple rearing and multiplica-
tion of queens, where individual queens
are evaluated predominantly by their egg
laying ability and body size. ose two
markers of queen quality (fecundity and
size) are certainly important for the prop-
agation of queens, but selective breeding
requires more than propagation. Selective
breeding implies the intentional selection
for genetic improvement of the population
as a whole, with every new generation
improved compared to the previous one,
ideally for all traits of interest.
To achieve such improved selection,
queens and colonies in each generation
must be chosen that exhibit the desirable
properties the breeder wishes to propa-
gate, using these as parents for the next
generation so that on average the next
generation is expected to be better than
the previous one. Breeding success for a
certain honey bee population is thus the
cumulative outcome of many actions,
such as performance testing on the colony,
the selection and mating of individuals.
Our aim with this paper is to help demys-
tify conventional honey bee breeding pro-
grams and highlight the typical features of
successful programs, based on the breed-
ing experiences in Central Europe. We
focus on conceptual aspects beekeepers
and breeders can control without discuss-
ing the underlying theories based upon
population and quantitative genetics.
A Breeding Program and
Its Elements
A breeding program represents a set of
systematically planned and implemented
activities aimed at the sustained genetic
improvement of a honey bee population
(Brascamp, 2014; Tiesler et al., 2016).
us, by continuous implementation of
this selection program it is expected that
the colonies in the next generation will
express improved behavior concerning
targeted traits (for instance: gentleness,
calmness on the comb, reduced swarm-
ing), enhanced production (honey, pollen,
wax, royal jelly) and vitality (resistance to
diseases and pests, prolonged life expec-
tancy, etc.).
A breeding program should include
explicit breeding objectives, performance
testing to evaluate the desired character-
istics, estimation of the breeding values,
selection, mating, multiplication of the
improved genetic stock, and evaluation
(Figure 1).
e rst and very important step is to
dene the breeding objective. us, based
on the economic importance, scientic
evidence and practical experience, breed-
ers must decide which traits they intend to
improve, and what is the relative impor-
tance of improving the dierent traits.
Generally, the preferred traits in selection
for honey bees involve improving honey
yield, gentleness, decreasing the swarming
tendancy and increasing Var r o a resist-
ance. Sometimes traits such as coloration,
which are typical for a particular popula-
tion, can be included in the selection pro-
cess. In principle all traits of importance
should be included, because selection
for too limited a set of traits may lead to
deterioration of other relevant traits. For
example selecting only for Var ro a resist-
ance with no regard for the qualities that
make for productive colonies can result in
queens that are not practical for commer-
cial beekeeping.
e German breeding program for
resistance to Var r o a (AGT) is an example
where the focus is on Var ro a resistance
(Table 1), but other traits are also taken
into account. e decision regarding
which traits to include to achieve the
breeding objective depends mainly on the
interests of a particular group of breeders.
Objectives such as the conservation of an
isolated population or specic research
goals can be relevant too.
Since breeding success is accumulated
genetic improvement over time the
© 2017 International Bee Research Association
Figure 1. Elements of a breeding program and sequence of actions. A continual process
of selecting towards the breeding objective. Mating is carried out to produce a next
generation for performance testing, but also for multiplication of the improved material to
be disseminated and utilized in the broad population.
DOI: 10.1080/0005772X.2017.1345427
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© 2017 International Bee Research Association
breeding objective ideally should be con-
sistent from year to year and not subject
to frequent drastic changes. us, before
the objective is dened one should foresee
the needs and demands that might occur
in the future, so that the selection has
long-term relevance. Finally, the breeding
objective should take into account the way
bees are managed and how beekeeping
operations are organized if the goal is
improved stock for beekeepers.
Performance testing requires a standard-
ized methodology for the phenotypic
assessment of traits and accurate data
recording (Büchler et al., 2013; Ruttner,
1972; Uzunov et al., 2015). While in farm
animals performance is generally meas-
ured on individuals (milk production in
dairy cows, egg laying rate in hens), in
honey bees traits are usually observed
on the colony level. is is illustrated in
Figure 2.
On the one hand the workers contribute
to colony’s performance, known as the
worker eect, but the queen (1a) also has
a direct eect (queen eect) on the colo-
ny’s performance. Of course she also con-
tributes indirectly, because she transmits
her breeding value for the worker eect to
the workers. As an example, honey yield is
aected by the ability of workers to collect
nectar, but also by the egg laying capacity
and pheromone production of the queen.
Figure 2 also illustrates that apart from
the queen (1a) that is the mother of the
colony, the drone-producing queens (1b)
also contribute 50% to the breeding value
of the workers. Since drones are haploid
and have no father, it is oen said that
the paternal contributer to the breeding
value of the workers in practice is 4a, the
so-called father colony.
A written guideline for performance
testing (protocol), along with practice
and training of the bee breeders in the
proposed methods (breeders’ capacity
building) are essential so that test colonies
can be evaluated objectively and accu-
rately across apiaries and operations.
Before engaging in a breeding program
and queen evaluation, a number of pre-
paratory activities should be conducted.
ese include establishing test apiaries
and colonies, as well as organizing a
queen exchange among the dierent test
apiaries.1
e next step is the estimation of breeding
values. By denition the breeding value
of an individual is the expected perfor-
mance of the ospring when the indi-
vidual is mated to an average mate and
the ospring is performing in an average
environment. Usually these compli-
cated calculations of the breeding value
estimation are conducted by the facility
that manages the database of performance
testing results.
A best case study of such a system is
represented by BEEBREED (www.
beebreed.eu) where via an online web-
based platform the performance testing
data are stored, breeding values are
estimated and subsequently published. In
BEEBREED the estimated breeding value
of an individual takes into account the
colony’s individual performance as well as
the performances of other colonies in the
same environment (test apiary) and per-
formances of colonies that are ancestors
or other relatives (Figure 3). Comparison
of colony’s performance with that of
colonies at the same test apiary takes into
account dierences caused by beekeeping
techniques applied, weather, food sources
etc. For more details, see Bienefeld et al.
(2007), Büchler et al. (2013), Brascamp
et al. (2016) and Tiesler et al. (2016).
Table 1. The weighting of desirable traits in the AGT breeding program.
aA combination of natural mite fall in spring, mite infestation in late summer and hygienic behavior.
Trait Honey yield Gentleness Calmness Swarming behavior Varroa indexa
Weighting of traits (%) in the total breeding value 15 15 15 15 40
Figure 2. Effects of the workers and queen
on the colony’s performance.
Figure 3. Simplied model of the effects contributing to the estimation of breeding
values according to BLUP Animal Model.
DOI: 10.1080/0005772X.2017.1345427
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To obtain reliable breeding values, it is
important that daughters of each breeder
queen are tested at several test apiaries.
e selection of queens based upon
ranking for estimated breeding values is
a further step in the breeding program.
Usually individual breeders not only select
based upon this criterion but include
additional observations such as overwin-
tering ability, colony strength, etc. us
queens are ranked and selected as mothers
for the next generation of queens as well
as queens to produce drone producing
queens. Oen individual breeders decide
what queens they will use to produce the
next generation, while an association is
oen responsible for selecting the queen
that will be the mother of the queens
that head the multiple drone colonies at
a mating station, under the assumption
that such a mating station is not used
by one single breeder. Via instrumental
insemination or private isolated mating
yards, an individual breeder can control
which queen stock serves as the paternal
ancestor of the next generation.
In honey bees mating deserves specic
attention due to the reproductive behav-
ior of the species. Special measures are
required to control mating, for exam-
ple through the use of mating stations
(isolated mainland or island locations) or
instrumental insemination. According to
theory the breeding value of a progeny
equals half the sum of the breeding values
of the parents. As a consequence, parents
can be selected independently and mated
at random when it concerns the average
breeding value of the next generation.
However, the desire of individual breed-
ers to compensate for a weakness in a
selected queen with the strength of the
mate advocates for the combined selection
of a queen and her mates and not just the
individual parents. For example a queen
that has excellent Var r o a resistance and
honey production, but a higher swarm
tendency might be mated with drones that
originated from a queen that exhibited a
very low swarm drive. Another benet of
selecting both partners is that potential
inbreeding of workers can be taken into
account.
Genetic improvement is the ultimate
goal of a breeding program and involves
eventual propagation of the improved
stock in the general honey bee population
(Büchler et al., 2010; Rinderer et al., 2010).
e beekeeping community can incorpo-
rate improved stock on the maternal side
in the form of graed larvae, queens cells,
virgin or mated queens. rough the use
of drone semen in instrumental insemina-
tion or the mating stations where selected
queens produce drones that will mate the
virgin queens, beekeepers can incorporate
improved genetic stock on the paternal
side, helping to improve the quality of the
bee population.
We feel that the topic of propagation
of improved stock does not receive the
attention it deserves. Activities concerning
this goal, like transparent communication,
marketing and ecient delivery systems,
should be well planned and implemented
and require concerted actions of the bee-
keeping community at large.
Finally, a program should systematically
be subjected to evaluation. is should
be recognized as a distinct element. e
rst step is to compare the predicted
and realized selection response or “does
the program meet its promises?”. In the
evaluation, all elements and steps of the
breeding program should be scrutinized
and any obstacles should be identied and
removed.
Here issues like the accuracy of the meth-
odology for data gathering, the recording
of pedigrees, and the accuracy of breeding
value estimations should be evaluated, as
well as re-evaluating strategic goals and
delineating future expectations. In addi-
tion, breeding programs should consider
if the environments in which the daughter
queens of the breeding program perform
are suciently uniform, such that the col-
onies produced from the better line in one
environment are consistently the better
ones in another environment.
Types of Breeding
Programs
In the previous paragraphs we described
a breeding program based strongly on the
results of BEEBREED and the beekeeping
community in Europe. However, the rea-
sons to keep, and possibly improve honey
bees, dier among bee breeders around
the world.
ere is a wide spectrum of breeding pro-
grams. To illustrate this we list the most
common types of breeding programs.
• Commercial, aimed to improve the
overall performance of the honey bees
from the population of interest, based
upon the assessment of various traits.
Occasionally, the number of traits is
limited to 3 or 4. However, the main
drive and objective of these commercial
breeding programs is improvement in
commercially important traits (more
honey, less defensive bees, reduced
swarming tendency, etc.). is type of
selection is most common and consid-
ered by us as the most sustainable.
• Conservation, aimed at the maintenance
of endangered honey bee populations.
e ultimate goal is maintenance or
enlargement of the population. Genetic
improvement of such populations is a
useful tool in the context of “conser-
vation by utilization”, as conservation
by utilization is considered a preferred
mechanism to conserve subspecies or
populations. Along with commonly rec-
ognized traits, relevant on the regional
or local scale, morphological characters
and molecular markers are frequently
the basis for decision-making and selec-
tion, the latter two being used to ensure
that the population is not mixed with
other subspecies.
• Research breeding programs can be ini-
tiated for studying certain traits (eects
of the genes, identication of markers,
etc.) of scientic interest as well as
analyzing the eects of hybridization or
inbreeding, assessing the adaptive abil-
ity of populations, resistance to diseases,
genotype by environment interactions,
etc. Generally these breeding programs
are short-term and under the respon-
sibility of research institutes or other
academic institutions.
Whatever type of breeding program is
initialized there is always the question
of the genetic origin of the population
under selection. In general there are two
main approaches. Recently a trend has
emerged for sustainable conservation, a
popular breeding approach to improve
and conserve the native or locally adapted
honey bee populations or subspecies. e
basic philosophy behind this is to reduce
importations and instead utilize and
improve the local populations in compari-
son to the non-local ones.
Alternatively, oen breeders start a breed-
ing program building upon breeds like
Ligustica, Carnica or Buckfast (which can
be considered a breed of recent origin)
with assumed general usefulness accross
environments. From a conservation point
of view the disadvantage of this is that
the mixing of local populations and these
general purpose populations, when kept
in the same area, is dicult to prevent.
Whatever approach is chosen the main
issue during the initiation of the breeding
program is appropriate selection of the
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Bee World • VOL 94 • September 2017 • Page 87
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rst genotypes. Here the best advice is to
choose the best available queens, prefer-
ably based on a breeder’s records, taking
care that the founding queens represent
sucient genetic diversity.
Conducting the Breeding
Program
Breeders, regional groups, scientists,
national and local authorities are the
main partners of a breeding program.
e organization of regular meetings is
important for better coordination of the
activities as well as for exchange of ideas
and overcoming the challenges of imple-
mentation. A close cooperation with sci-
entists can signicantly help the program
achieve its breeding objective, by aiding in
the development of standardized meth-
odology and introducing new approaches
and models of breeding.
Concerning the establishment of con-
trolled mating locations, the cooperation
with local authorities may be important.
If we take in to consideration the above
mentioned elements, the overall success
and sustainability of a breeding program
signicantly depends on the collabora-
tion, communication, transparency and
exchange of ideas among all stakeholders
within and around the breeding program.
Acknowledgment
We would like to thank Dr. Bjorn Dahle,
Dr. Sreten Andonov, Dr. Eliza Cauia, Egoitz
Galarza, Borce Pavlov, Dr. Andreas Hoppe
and Dirk Ahrens-Lagast for their valuble
comments and suggestions for improvement
of the readability of the article and integra-
tion of experience from various breeding
practices across Europe.
Note
1. Location or apiary where testing of the colonies is
performed.
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A. Uzunov and R. Büchler
Bee Institute in Kirchhain, Erlenstrasse 9,
Kirchhain 35274, Germany
Email: aleksandar.uzunov@llh.hessen.de
E. W. Brascamp
Wageningen University & Research
Animal Breeding and Genomics,
P.O. Box 338, 6700 AH, Wageningen, e
Netherlands
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