Journal of Apicultural Research

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The effect of surface area of pollen patties fed

to honey bee (Apis mellifera) colonies on their
consumption, brood production and honey yields

Dorit Avni, Arnon Dag & Sharoni Shafir

To cite this article: Dorit Avni, Arnon Dag & Sharoni Shafir (2009) The effect of surface area
of pollen patties fed to honey bee (Apis mellifera) colonies on their consumption, brood
production and honey yields, Journal of Apicultural Research, 48:1, 23-28

To link to this article: http://dx.doi.org/10.3896/IBRA.1.48.1.06

Published online: 02 Apr 2015.

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 Journal of Apicultural Research and Bee World 48(1): 23-28 (2009) © IBRA 2009
DOI 10.3896/IBRA.1.48.1.06

ORIGINAL RESEARCH ARTICLE

The effect of surface area of pollen patties fed to

honey bee (Apis mellifera) colonies on their
consumption, brood production and honey yields.

Dorit Avni¹, Arnon Dag² and Sharoni Shafir¹*.

¹B. Triwaks Bee Research Center, Department of Entomology, Faculty of Agricultural, Food & Environmental Quality Sciences,
The Hebrew University of Jerusalem, Rehovot, 76100, Israel.
²Gilat Research Center, Institute of Plant Sciences, Agricultural Research Organization, 85280, Israel.

Received 6 January 2008, accepted subject to revision 14 May 2008, accepted for publication 17 August 2008.
Downloaded by [University of California, San Diego] at 09:45 11 February 2016

*Corresponding author: Email: shafir@agri.huji.ac.il

Summary
Beekeepers may feed colonies with pollen supplement patties during times of pollen shortage. We tested whether the surface area of pollen
patties fed to honey bee colonies affects their consumption, brood production and honey yields. We compared three pollen-patty sizes, of
equal weight, and found that consumption increased with surface area. Consequently, brood production tended to increase with pollen-patty
size, and colonies fed the patty with the largest surface area produced significantly more brood than those fed a carbohydrate-only control
patty. The colonies in the large surface area treatment also tended to produce more honey, though the differences were not statistically
significant, probably due to the time gap between the pollen deficiency period and the time honey was harvested. We conclude that pollen
patties with greater surface area are more readily consumed by honey bee colonies and may contribute more to colony development and
subsequent honey yields.

Efecto de la superficie de área de la tarta de polen en colonias de la

abeja melífera (Apis mellifera) en su consumo, producción de cría
y de miel.
Resumen
Los apicultores pueden alimentar a las colonias de abejas con tartas de suplemento de polen durante épocas de escasez de este producto.
Probamos si la superficie de área expuesta de la tarta de polen afecta su consumo, producción de cría y de miel en la colonia de abejas.
Comparamos tres tamaños de la tarta de polen, con el mismo peso, y encontramos que el consumo incrementó con la superficie expuesta.
En consecuencia, la producción de cría tuvo una tendencia al incremento con respecto al tamaño de la tarta de polen, las colonias alimentadas
con la tarta con mayor superficie expuesta produjeron significativamente más cría que aquellas alimentadas con la tarta control a base de
carbohidrato únicamente. Las colonias con el tratamiento de mayor superficie expuesta también tendieron a producir más miel, aunque las
diferencias no fueron estadísticamente significativas, probablemente debido al espacio de tiempo entre el período de escasez de polen y el
tiempo en que la miel fue cosechada. Concluimos que las tartas de polen con mayor superficie de exposición fueron consumidas con mayor
facilidad por las colonias y pueden contribuir mayormente al desarrollo de la colonia y en la producción de miel.

Keywords: pollen patty size, consumption, Apis mellifera, brood production, honey yield.
 24
Introduction
Honey bees feed on nectar and pollen, which provide them the
carbohydrates, proteins, lipids, vitamins and minerals that are
required for their proper development (Winston, 1987; Cohen, 2004;
Keller et al., 2005a). Nectar is their main source of energy, and
pollen is necessary for reproduction and brood rearing (Dietz, 1978;
Herbert, 1999). Honey bee colonies may be exposed to periods of
shortage of one or more of their nutritional demands. For example, in
some locations or seasons there may be a shortage of nectar,
causing bees to use their honey stores (Day et al., 1990; Keller et al.,
2005b). There may also be a shortage of pollen, and bees then re-
duce the amount of brood that they rear (Day et al., 1990). Colonies
used for crop pollination in enclosures, such as greenhouses, net
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houses, and net covered orchards, may often experience nutritional
deficits (Kalev et al., 2002; Keller et al., 2005b). Under such
conditions, bees are limited to one type of crop, which might lack, or
contain only small amounts, of nectar and pollen (Day et al., 1990).
Honey bees can deal with periods of shortage in nectar and
pollen within their foraging range, by lowering their metabolism and
reducing tasks and activities in the hive, such as foraging and brood
rearing (Imdorf et al., 1998; Pernal and Currie, 2001; Kalev et al.,
2002; Keller et al., 2005b). Less brood rearing eventually reduces the
number of adult bees, including foragers, and may consequently
affect pollination efficiency and honey yields (Duff and Furgala, 1986;
Nelson, 1987; Fewell and Winston, 1992; Herbert, 1999).
If nectar is insufficient, beekeepers can supply the bees with
sugar syrup, which is a routine beekeeping practice. If pollen is
lacking, beekeepers may use pollen substitutes or supplements in
order to strengthen the colony. A wide range of plant and animal
protein sources have been tested as substitutes for pollen, including
soybean flour, peanut meal, whole wheat flour, brewers yeast, corn
flour, ground dry fish, powdered skim milk, egg albumin, and others
(Erickson and Herbert, 1980; Doull, 1980; Lehner, 1983; Kalev et al.,
2002; Keller et al., 2005b). Supplements contain bee-collected pollen
mixed with other ingredients, for example soybean flour and honey,
in order to form the desired patty consistency (Kalev et al., 2002;
Keller et al., 2005b). Patty composition is important both for its
nutritional value and for its effect on how readily bees consume it
(Herbert, 1999; Keller et al., 2005). Bees tend to consume pollen
supplements more readily than pollen substitutes (Standifer et al.,
1973), and as a result of the higher consumption, to rear more brood
(Keller et al., 2005b).
Whereas much research has focused on pollen patty
composition, the effect of how the patty is applied has received less
attention. Pollen-patty application method can influence the level of
patty’s consumption and therefore its effectiveness (Cook and
Wilkinson, 1986; Herbert, 1999; Keller et al., 2005a, 2005b). The
Avni, Dag, Shafir
present study for the first time tests the effect of surface area of a
pollen patty on its consumption by honey bees, on brood production
and on subsequent honey yields.
Materials and methods
The feeding phase of the experiment was conducted between De-
cember 2004 and January 2005 in Moshav Tal–Shahar (31°48′00′′ N,
34°54′60′′ E, elevation 141m) in the Judean Hills, central-eastern
part of Israel. Pollen sources are usually limited during the winter in
this location.
Bee colonies and assessments
On 6 December 2004, we selected 40 hives, each with 7-10 popu-
lated frames and 4-6 brood frames. The queens were three-months
old, daughters of the same mother queen from the local breeding
programme of the Ministry of Agriculture apiary in Zriffin, which is
based on the Italian strain. We assessed the hives following a stan-
dard procedure that included an estimate of the number of populated
frames, and the total comb area with sealed brood, open brood
(eggs and larvae), stored pollen, and stored honey. Area was esti-
mated by dividing each side of each comb into eight squares (100mm
X 100mm = 1dm²), and counting the number of such squares of
each type (Kalev et al., 2002). We then divided the hives into four
treatment groups in a pseudorandom fashion. Colonies were first
divided into ten groups having similar means and variances of sealed
brood area. Then each group was randomly assigned to one of the
four treatments. We continued to assess the hives every three
weeks, for three more assessments.
Treatments and feeding
We fed patties to the colonies every 10 days, from 6 December until
4 February 2005, when pollen from the local flora became abundant
and the bees started storing pollen in the hives. There were four
treatments, three pollen supplement patties that differed in size, and
a carbohydrate only control of medium size. All patties weighed 300g
each, but differed in surface area. Small, medium, and large patties
had dimensions of 100 x 100 x 23 mm, 141 x 141 x 11.5 mm and
200 x 200 x 5.8 mm, respectively. The surface areas of the three
sizes were 29,200 mm2, 46,248 mm2 and 84,640 mm2, respectively.
The pollen supplement patties contained 26.7 % roasted soy flour
(48 % protein and <2 % lipids; “Tavliney Hagalil” Ltd.; Afula), 26.7
% bee-collected pollen, 26.7 % sugar (sucrose) powder (“Oz asakim”
Ltd.; Ashdod) and 19.9 % honey. The control patty contained 67 %
sugar powder and 33 % honey. Patty composition was based on
Kalev et al. (2002).
 Pollen patty surface area and consumption 25

At each feeding, we placed two patties (600 g) on a plastic small medium large control
30
grid above the brood area in each hive. The grid had openings of

Pollen stores (sqr

25
6.5 mm, which allowed bees access to the patty from below. Material

decimeters)
20
left over from the previous feeding was collected and weighed.
15

10
Honey yields
5
In late February we moved the hives to an area near a large forest of
0
flowering Eucalyptus trees. Two months later we extracted the honey 6.12.04 27.12.04 26.1.05 16.2.05

from the hives; honey supers from each hive were weighed before Dat e

and after extraction, thus providing net weight of extracted honey

Fig. 2. Mean pollen stores (± s.e.) in hives fed pollen patties of three
per colony.
surface areas and a sugar control between December 2004 and
February 2005. There were no statistically significant differences
Data analysis
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between treatments in any sampling date.

Of the original 40 colonies, six died or were excluded from the
experiment because of queen replacement or theft throughout the
experiment period. These excluded colonies were equally distributed
Results
between treatments, with one or two colonies excluded from each
treatment. One more colony was stolen before its honey was extracted. Consumption
Bees consistently consumed the control patty most, followed by
small medium large candy pollen supplement patties from largest to smallest surface area (Fig.
650 1). The effect of patty size on consumption levels varied between
a a a ab a ab a a a
sampling dates (Time x treatment: F9.9, 98.7 = 2.33, P = 0.017).
Consumption (grams )

600 ab bc
b b
c
550 bc
bc However, consumption of the large patties was significantly greater
b b
c than that of the small ones in most of the cases (Fig. 1).
500 bc b c
bb
450
Pollen stores
400 c
Pollen stores were relatively low in all treatments during the first
350
seven weeks of the experiment, and rose dramatically (up to six-fold)
300
by the last assessment (Fig. 2). The effect of patty size on pollen
17.12.04 27.12.04 7.1.05 17.1.05 26.1.05 4.2.05

Dat e stores varied between sampling dates (Time x treatment: F8.3, 82.7 =
2.12, P = 0.041) However, pollen stores were generally similar
Fig. 1. Mean (± s.e.) consumption of pollen patties of three surface
areas and a sugar control between December 2004 and February
2005. Different letters in each measuring date represent means that
are statistically different (Tukey test, P<0.05). small medium large control
40
a a
35 a
a
We used repeated measures of analysis of variance to a a
30
a
Sealed brood (sqr

compare between treatments over the sampling dates. The reported 25 a

decimeters)

probability values are based on the conservative correction according 20

a a a
to the univariate adjusted Geisser-Greenhouse degrees of freedom a a
15 a a
(JMP v.7, SAS Institute, Inc., Cary, NC). To compare between 10
b
treatments within sampling dates we performed multiple comparison 5
post tests using Tukey’s method, at a probability level of 0.05. Sealed 0
brood data were transformed by the square-root transformation to 6.12.04 27.12.04 26.1.05 16.2.05
meet ANOVA requirements. The consumption and pollen storage data Date
did not conform to requirements of a parametric ANOVA, so we used
a nonparametric ANOVA on ranks (Conover and Iman 1981). Honey Fig. 3. Mean (± s.e.) sealed brood areas in hives fed pollen patties
yields were compared between treatments using ANOVA. Means are of three surface areas and a sugar control between December 2004
reported with standard errors. and February 2005. Different letters represent means that are
statistically different (Tukey test, P<0.05).
 26 Avni, Dag, Shafir

2 a Honey yields
Difference in sealed brood area

0 There were no statistically significant differences in honey yields

between treatments (F3,29=1.21, P=0.32). However, colonies that
(sqr decimeters)

-2

ab
received the large pollen supplement patties tended to produce more
-4 ab
honey (31.3 ± 3.5 kg) compared to those receiving the medium
small
-6 (24.3 ± 3.6 kg) and small (24.8 ± 3.0 kg) patties. Interestingly,
medium
-8
honey yields were also relatively high in the control group (30.0 ±
large
control
b 2.8 kg).
-10
Treatment

Fig. 4. Change in sealed brood area in hives fed pollen patties of

Discussion
three surface areas and a sugar control during extreme pollen Absence of pollen may affect the strength of the colony and its honey
shortage, between assessments on December 6 and December 27, production (Duff and Furgala, 1986; Nelson, 1987; Keller et al.,
Downloaded by [University of California, San Diego] at 09:45 11 February 2016

2004. Different letters represent means that are statistically different 2005a, 2005b). Beekeepers can feed colonies with pollen substitutes
(Tukey test, P<0.05). or supplements, but in order to be effective these must be eagerly
consumed by the colony (Standifer et al., 1973; Schmidt and Hanna,

between treatments during each assessment, and there was no 2006). Our results show that the patty surface area affects its con-

statistically significant difference between treatments during any of sumption levels.

the assessments when compared separately by Tukey’s method Colonies in this study experienced pollen shortage until mid

(Fig.2). February, when pollen stores increased about five fold (Fig. 2). The
most severe shortage was in December, as seen by the decline in
sealed brood of the control treatment between our first and second
Sealed brood
colony assessments (Fig. 3). Pollen probably started becoming avail-
The effect of patty size on area of sealed brood varied between
able in early January, since by our third assessment at the end of
sampling dates (Time x treatment: F8.1, 81.1 = 3.21, P = 0.003). On
January the control colonies had already greatly increased their
the second assessment, three weeks after the beginning of feeding,
brood production. Pollen brought into the colony after a period of
only the control treatment showed a significant drop in sealed brood
severe shortage was probably immediately consumed, leading to an
area (Fig. 3). There was a tendency during the second and third
increase in sealed brood in all treatments by late January. Pollen
assessments for sealed brood area to be greater in the colonies
stores during this time were still low, but were especially low in colo-
receiving the larger pollen patties. By the third assessment, brood
nies receiving the large patty compared to those receiving the small
areas had increased in all treatments, including the control. The
patty. The former may have consumed more of the incoming pollen,
roughly 4.5 fold increase in sealed brood in the control shows that
because they tended to have more brood. An alternative hypothesis
the main pollen shortage had ended by that time. We compared the
is that because the large surface patty was so effective, the bees
effect of the treatments during the period of greatest pollen short-
were more content with the pollen they got from the patty, and
age, between the first and second assessments (Fig. 4). Colonies
therefore foraged less for pollen. Overall, the amount of collected
from different treatments were affected differently by the pollen
and stored pollen is influenced by the dynamics of various factors in
shortage (F3,30=3.08; P=0.043). Only colonies fed the pollen patty
the colony, such as colony size, intrinsic bee characteristics and re-
with the greatest surface area did not suffer a decrease in sealed
source condition, and other factors in the environment, such as rain,
brood during this time, and colonies fed the control patty suffered
temperature and botanical resource availability (Hrassnigg and Crail-
the greatest decline in sealed brood.
sheim, 1998; Dimou et al., 2006; Schmickl and Crailsheim, 2007).
During the entire study, bees consumed more from the large
Honey stores
surface area pollen-patties than from the small ones. Interestingly,
Patty size did not affect honey stores (Treatment: F3, 30 = 0.28,
the control patties, which were composed only of carbohydrates,
P = 0.84) throughout the experiment (Time x treatment: F5.1,50.9
were consumed the fastest. This may be due to a phagostimulating
= 1.37, P = 0.25). However, mean honey stores declined from 89.7
effect of the high sucrose content, since honey bees are highly re-
square decimeters during the first assessment to 48.1 square
sponsive to sucrose (Scheiner et al., 2004) and phagostimulants
decimeters during the last assessment (Time F1.7, 50.9 = 191,
greatly affect patty consumption (Keller et al., 2005a, 2005b;
P < 0.0001).
Schmidt and Hanna, 2006). In addition, bees store carbohydrates in
the comb cells, as they store nectar, but apparently consume pollen
 Pollen patty surface area and consumption
substitutes or supplements directly and do not store them in the
comb cells as bee bread (Crailsheim et al., 2006). Despite their high
consumption, control patties could not sustain brood during a pollen
dearth period. Colonies fed pollen supplements typically maintain
higher brood levels than those fed a carbohydrate only diet
(Standifer et al., 1973; Doull, 1975; Doull, 1980; Cook and Wilkinson,
1986; Imdorf et al., 1998).
Feeding pollen supplements, under pollen shortage
conditions, can lead to higher honey yields (Doull, 1980; Nabors,
2000). We did not find significant differences between treatments in
the amount of honey we extracted from the colonies in late April. The
severe pollen shortage was only in December, and differences in
sealed brood between treatments disappeared in January with the
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influx of pollen. Between then and April, colonies reached similar
strength, and consequently similar honey yields, even though there
was a tendency for the colonies to reach higher honey yields when
they received larger surface area patties during the dearth period.
Whereas the control colonies appeared to have relatively high honey
yields, some of it may have been derived from the sugar patties that
they were fed rather than from floral nectar. The advantages of a
superior diet would manifest themselves more the shorter the time
between a pollen dearth and the honey flow. This statement is
supported by the finding of Slabezki et al. (2001) that feeding with
pollen supplements increased the yield of honey harvested one
month after the feeding period, but not five months later.
In conclusion, pollen substitute and supplement patties can
sustain colonies during periods of pollen shortage, leading to
stronger colonies for pollination and honey production. It is well
established that patty composition can affect its consumption and its
nutritional value. We show that the surface area of the patty also
affects its consumption and its contribution to brood rearing. In
particular, a patty with greater surface area is consumed more
quickly, and contributes to stronger colonies.
Acknowledgements
This study was funded by MERC Grant No. TA-MOU-03-M22-023. We
wish to thank Haim Kalev, Orit Rot, Erez Tsur and Holy Robaz for
their technical assistance.
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