cGMP-dependent changes in phototaxis: a possible role for the foraging gene in honey bee division of labor
Y. Ben-Shahar1,*,
H.-T. Leung3,
W. L. Pak3,
M. B. Sokolowski4 and
G. E. Robinson1,2
1 Department of Entomology, University of Illinois at Urbana-Champaign, 320
Morrill Hall, 505 S. Goodwin Avenue, Urbana, IL 61801, USA
2 Neuroscience Program, University of Illinois at Urbana-Champaign, 320
Morrill Hall, 505 S. Goodwin Avenue, Urbana, IL 61801, USA
3 Department of Biological Sciences, Purdue University, West Lafayette, IN
47907, USA
4 Department of Zoology, Mississauga Campus, University of Toronto, 3359
Mississauga Road, Mississauga, ON L5L1C6, Canada

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Fig. 1. Phototaxis apparatus. The proportion of bees that walked from the cage to
the light in 3 min was used as an index of positive phototaxis.
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Fig. 4. Electroretinogram (ERG) measurements in honey bees as a function of
behavioral state or treatment. N=6 bees tested at each light
intensity, per group. There were no significant differences between the groups
(P>0.05; repeated-measures ANOVA).
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Fig. 2. Behaviorally related differences in Amfor brain expression. Bars
represent relative mRNA differences, normalized to nurses (N). Error bars
represent the standard errors of the Ct converted to the same arbitrary
scale as the means (see Materials and methods). P values on graphs
refer to ANOVA; different letters denote statistically significant differences
(P<0.05; pair-wise LSD post hoc analysis).
N=810 brains per group (see numbers on bars), each brain was
analyzed individually. FH, food handlers; U, undertakers; F, foragers.
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Fig. 3. Phototaxis behavior in honey bees as a function of behavioral state or
treatment. (A) Differences in positive phototaxis between nurses and foragers.
Group size=20 bees. 2x2 2 tests were conducted
for both trials. (B) Effects of 8-Br-cGMP, 8-Br-cAMP on positive phototaxis.
Nine trials were conducted; data on the proportion showing positive phototaxis
from each trial were arcsine transformed for ANOVA. For graphical purposes,
means ± S.E.M. were back-transformed (resulting in
asymetrical error bars). For doses, see Materials and methods. `Light',
phototaxis behavior when light source is on; `Dark', locomotion control when
light source is off.
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Fig. 5. Effects of 8-Br-cGMP on levels of locomotor activity, age at onset of
circadian locomotor rhythms, and tau (the period of rhythmicity)
under (A) 12 h:12 h light:dark (L:D) and (B) 24 h dark (D:D) regimes. Activity
levels were analyzed only under L:D conditions because light entrains
activity. There were no significant differences between the different groups
in all experiments (activity, tau, ANOVA; % rhythmicity, P>0.05;
2 tests).
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Fig. 6. Brain mRNA levels for genes encoding PKA regulatory (r) and catalytic (c)
subunits as a function of behavior. Analysis and graphic representation are as
in Fig. 2. cDNA samples were
the same as in Fig. 2, trial 1
(with same sample sizes).
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© The Company of Biologists Ltd 2003