T1R2+T1R3-independent chemosensory inputs contributing to behavioral discrimination of sugars in mice.

Simple sugars are thought to elicit a unitary sensation, principally via the "sweet" taste receptor type 1 taste receptor (T1R)2+T1R3, yet we previously found that rats with experience consuming two metabolically distinct sugars, glucose and fructose, subsequently licked more for glucose than fructose, even when postingestive influences were abated. The results pointed to the existence of an orosensory receptor that binds one sugar but not the other and whose signal is channeled into neural circuits that motivate ingestion. Here we sought to determine the chemosensory nature of this signal. First, we assessed whether T1R2 and/or T1R3 are necessary to acquire this behavioral discrimination, replicating our rat study in T1R2+T1R3 double-knockout (KO) mice and their wild-type counterparts as well as in two common mouse strains that vary in their sensitivity to sweeteners [C57BL/6 (B6) and 129X1/SvJ (129)]. These studies showed that extensive exposure to multiple concentrations of glucose and fructose in daily one-bottle 30-min sessions enhanced lick responses for glucose over fructose in brief-access tests. This was true even for KO mice that lacked the canonical "sweet" taste receptor. Surgical disconnection of olfactory inputs to the forebrain (bulbotomy) in B6 mice severely disrupted the ability to express this experience-dependent sugar discrimination. Importantly, these bulbotomized B6 mice exhibited severely blunted responsiveness to both sugars relative to water in brief-access lick tests, despite the fact that they have intact T1R2+T1R3 receptors. The results highlight the importance of other sources of chemosensory and postingestive inputs in shaping and maintaining "hardwired" responses to sugar.

Navigation.

Experiments with migrating birds displaced during autumn migration outside their normal migration corridor reveal two different navigational strategies: adult migrants compensate for the displacement, and head towards their traditional wintering areas, whereas young first-time migrants continue in their migratory direction. Young birds are guided to their still unknown goal by a genetically coded migration program that indicates duration and direction(s) of the migratory flight by controlling the amount of migratory restlessness and the compass course(s) with respect to the geomagnetic field and celestial rotation. Adult migrants that have already wintered and are familiar with the goal area approach the goal by true navigation, specifically heading towards it and changing their course correspondingly after displacement. During their first journey, young birds experience the distribution of potential navigational factors en route and in their winter home, which allows them to truly navigate on their next migrations. The navigational factors used appear to include magnetic intensity as a component in their multi-modal navigational 'map'; olfactory input is also involved, even if it is not yet entirely clear in what way. The mechanisms of migratory birds for true navigation over long distances appear to be in principle similar to those discussed for by homing pigeons.

Considerations on the role of olfactory input in avian navigation.

A large amount of data documents an important role of olfactory input in pigeon navigation, but the nature of this role is not entirely clear. The olfactory navigation hypothesis assumes that odors are carrying essential navigational information, yet some recent experiments support an activating role of odors. This led to an ongoing controversy. An important, often-neglected aspect of the findings on olfaction is that olfactory deprivation affects avian navigation only at unfamiliar sites. The orientation of anosmic birds at familiar sites remains an enigma; earlier assumptions that they would rely on familiar landmarks have been disproven by the home-oriented behavior of anosmic pigeons additionally deprived of object vision, which clearly indicated the use by the birds of non-visual, non-olfactory cues. However, if odors activate the establishing and enlarging of the navigational 'map' and promote the integration of local values of navigational factors into this map, it seems possible that such a process needs to occur only once at a given site, when the birds are visiting this site for the first time. If that were the case, the birds could interpret the local factors correctly at any later visit and orient by them. This hypothesis could explain the oriented behavior of birds at familiar sites, and it could also help to reconcile some of the seemingly controversial findings reported in the literature, where the effect of olfactory deprivation was reported to differ considerably between the various pigeon lofts, possibly because of different training procedures.

Mathematical analysis of the homing flights of pigeons based on GPS tracks.

To analyse the effect of magnetic and olfactory deprivation on the homing flight of pigeons, we released birds from a familiar site with either their upper beak or their nostrils anaesthetized. The tracks were analysed by time lag embedding to calculate the short-term correlation dimension, a variable that reflects the degrees of freedom and thus the number of factors involved in a system. We found that higher natural fluctuations in the earth's magnetic field characterized by A P-indices of 8 and above caused a reduction of the correlation dimension of the control birds. We thus separated the data into two groups according to whether they were recorded on magnetically quiet days or on days with higher magnetic fluctuations. Anaesthetizing the upper beak had no significant effect. Making pigeons anosmic reduced the correlation dimension on magnetically quiet days, but did not cause any reduction on days with higher fluctuations. Altogether, our data suggest an involvement of magnetic cues and olfactory factors during the homing flight and point to a robust, multi-factorial map.