Counterturns initiated by decrease in rate of increase of concentration

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Abstract
The position of beetles were marked at 1-sec intervals after they were released in still air 16–18 cm from point sources of pheromone. Characteristics of the tracks were quantified and compared to those that might be produced by counterturning schemakinesis, tropotaxis, klinotaxis, zigzagging, look-and-leap, or steepest-ascent schemakinesis mechanisms. The beetles' movements were highly irregular, but they turned almost continually and never fixed on a heading near 0° (=straight towards the source). Turn angle sizes increased slightly with absolute size of heading but had the opposite sign, thus compensating slightly for heading. Their distribution was centered about 0° and was unimodal. Heading decreased gradually as the source was neared, but the decrease became steeper within 1–5 cm of the source. Histograms showed that the maximum headings between occurrences when the beetle was headed directly towards the source (0°) were centered around 0° and most of them were less than 90°. However, maximum headings between 90° and 180° were not uncommon. Turn radius decreased as the source was neared. The counterturning mechanism was the most consistent with these observations. An analysis of rate of change of concentration with respect to heading and distance to the source further demonstrated that the counterturning mechanism could explain the form of the decrease in heading as the source was neared, if the major cue used to initiate counterturns was a decrease in the rate of increase of concentration. The tropotaxis could not recreate the form of the decrease, under any form of stimulus processing.