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First published online November 19, 2004
Journal of Experimental Biology 207, 4371-4381 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01306
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Floral scents induce recall of navigational and visual memories in honeybees

Judith Reinhard*, Mandyam V. Srinivasan, David Guez and Shaowu W. Zhang

Research School of Biological Sciences, Visual Sciences, The Australian National University, PO Box 475, Canberra, ACT 2601, Australia



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  		Fig. 1. (A) Design of feeders used during training and tests. (B) Setup of device used for blowing scent into the hive during the tests.

 


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  		Fig. 2. Experimental configuration, showing feeder locations relative to the hive for experiment 1 (A,B), experiments 2–4 (C), experiment 5 (D), and experiment 6 (E,F).

 


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  		Fig. 3. Experiment 1. Scent-triggered navigation of groups of bees trained to a single feeder. Left: test results for bees trained to a rose-scented feeder (white). Right: test results for bees trained to a lemon-scented feeder (grey). The figure shows means ± S.E. of visits by marked bees to an empty, unscented test feeder during an 8 min no-scent interval (air, unfilled bars) and an 8 min scent interval, when the respective scent is blown into the hive (hatched bars). N=9 tests for each scent and each no-scent interval. Numbers above bars show P-values for tests of significant difference between air and scent (Wilcoxon Matched-Pairs test).

 


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  		Fig. 4. Experiment 2. Scent-triggered navigation of groups of bees trained simultaneously to a rose-scented and a lemon-scented feeder placed at two different locations. The figure shows means ± S.E. of visits by marked bees to empty, unscented test feeders (white, rose feeder; grey, lemon feeder) during an 8 min no-scent interval (air, unfilled bars) and an 8 min scent interval (hatched bars), when either rose scent (above) or lemon scent (below) was blown into the hive. N=5 tests for each scent and each no-scent interval. Different letters above bars denote significant difference at a level of at least P<0.05 (Friedman ANOVA and Wilcoxon Matched-Pairs test).

 


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  		Fig. 5. Experiment 3. Scent-triggered navigation in individual bees, trained alternately to a rose-scented and a lemon-scented feeder placed at two different locations. Pie charts show distribution of visits to four empty, unscented test feeders during an 8 min rose scent interval, and an 8 min lemon scent interval (white, former rose feeder; grey, former lemon feeder; black, two dummy feeders placed randomly in the area). Data are accumulated from three tests for each scent, and are shown separately for first visits, circlings, landings and total visits. The number of individually marked bees that visited both test feeders, the number of choices (N) and P-values (observed vs expected frequency Chi2 Test) are shown below the pie charts. The expected frequency, based on random choice among the four test feeders, was 25%.

 


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  		Fig. 6. Experiment 3. Scent-triggered navigation in individual bees, trained alternately to two feeders, each placed in a different location and carrying a different scent. The figure shows the results of experiments using the following scent combinations: rose and lemon, rose and almond, and almond and lemon. Pie charts show distribution of visits to the two test feeders in each experiment, when the respective scents were blown into the hive (white, former rose feeder; light grey, former lemon feeder; dark grey, former almond feeder). In each experiment data are accumulated from four tests for each scent, and are shown separately for first visits, circlings, landings and total visits. The number of individually marked bees that visited both test feeders, the number of choices (N) and P-values for tests of significant difference between the two feeders when the respective scents were blown (Chi2 test) are shown below the pie charts.

 


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  		Fig. 7. Experiment 4. Scent-triggered navigation in individual bees, trained cyclically to a rose-scented, a lemon-scented, and an almond-scented feeder, each placed at a different location. Pie charts show distribution of visits to the three test feeders when the respective scents were blown into the hive (white, former rose feeder; light grey, former lemon feeder; dark grey, former almond feeder). In each experiment data are accumulated from four tests for each scent, and are shown separately for first visits, circlings, landings and total visits. The number of individually marked bees that visited all three test feeders, the number of choices (N) and P-values (observed vs expected frequency Chi2 Test) are shown below the pie charts. The expected frequency, based on random choice among the three test feeders, was 33%.

 


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  		Fig. 8. Experiment 5. Scent-triggered navigation in individual bees, trained alternately to a yellow rose-scented and a blue lemon-scented feeder. The positions of the feeders were randomly varied on the perimeter of a circular area (diameter 10 m), to train bees to the colours of the feeders rather than then positions. Pie charts show distribution of visits to two empty, unscented, but coloured test feeders when the respective scents were blown into the hive (yellow, yellow test feeder; blue, blue test feeder). (A) Test feeders were placed 10 m apart, at diametrically opposite positions on the circle perimeter. (B) Test feeders were placed next to each other. In each case, data are accumulated from four tests for each scent. For A results are shown separately for first visits, circlings, landings and total visits, while for B only total visits could be registered. The number of individually marked bees that visited both test feeders, the number of choices (N), and P-values for tests of significant difference between the two feeders when the respective scents were blown (Chi2 Test) are shown below the pie charts.

 

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© The Company of Biologists Ltd 2004