Francesca Odetti

Hippocampus and homing in pigeons: left and right hemispheric differences in navigational map learning

One‐month‐old, inexperienced homing pigeons, prior to any opportunity to learn a navigational map, were subjected to either right or left unilateral ablation of the hippocampal formation (HF). These pigeons were then held together with a group of age‐matched control birds in an outdoor aviary, where they were kept for about 3 months with the opportunity to learn a navigational map. When subsequently tested for navigational map learning at about 4 months of age posthatching, control and right HF‐ablated pigeons were equally good at orienting homeward from distant, unfamiliar locations, indicating successful navigational map learning. By contrast, left HF‐ablated pigeons were impaired in orienting homeward, indicating a failure to learn a navigational map. Interestingly, both right and left HF‐ablated pigeons displayed impaired homing performance relative to controls. These results suggest that different aspects of homing pigeon navigation may be lateralized to different hemispheres, and in particular, the HF of the different hemispheres. The left HF appears critical for navigational map learning, i.e. determining an approximate direction home from distant, unfamiliar locations. The right HF, and possibly the left HF as well, appear to play an important role in local navigation near the loft, which is likely based on familiar landmarks.

Functional asymmetry of left and right avian piriform cortex in homing pigeons' navigation

It has been shown that homing pigeons rely on olfactory cues to navigate over unfamiliar areas and that any kind of olfactory impairment produces a dramatic reduction of navigational performance from unfamiliar sites. The avian piriform cortex is the main projection field of olfactory bulbs and it is supposed to process olfactory information; not surprisingly bilateral lesions to this telencephalic region disrupt homing pigeon navigation. In the present study, we attempted to assess whether the left and right piriform cortex are differentially involved in the use of the olfactory navigational map. Therefore, we released from unfamiliar locations pigeons subjected, when adult, to unilateral ablation of the piriform cortex. After being released, the pigeons lesioned to the right piriform cortex orientated similarly to the intact controls. On the contrary, the left lesioned birds were significantly more scattered than controls, showing a crucial role of the left piriform cortex in processing the olfactory cues needed for determining the direction of displacement. However, both lesioned groups were significantly slower than controls in flying back to the home loft, showing that the integrity of both sides of the piriform cortex is necessary to accomplish the whole homing process.

Factors reducing the expected deflection in initial orientation in clock-shifted homing pigeons

SUMMARY To orient from familiar sites, homing pigeons can rely on both an olfactory map and visual familiar landmarks. The latter can in principle be used in two different ways: either within a topographical map exploited for piloting or in a so-called mosaic map associated with a compass bearing. One way to investigate the matter is to put the compass and the topographical information in conflict by releasing clock-shifted pigeons from familiar locations. Although the compass orientation is in general dominant over a piloting strategy, a stronger or weaker tendency to correct towards the home direction by clock-shifted pigeons released from very familiar sites has often been observed. To investigate which factors are involved in the reduction of the deviation due to clock-shift, we performed a series of releases with intact and anosmic pigeons from familiar sites in unshifted and clock-shifted conditions and a series of releases from the same sites with naive clock-shifted birds. Our data suggest that the following factors have a role in reducing deviation due to the clock-shift: familiarity with the release site, the lack of olfactory information and some unknown site-dependent features.

Bilateral participation of the hippocampus in familiar landmark navigation by homing pigeons

Recent findings indicate a different role of the left and right hippocampal formation (RHF) in homing pigeon navigational map learning. However, it remains uncertain whether the left or the RHF may play a more important role in navigation based on familiar landmarks. In the present study, we attempted to answer this question by experimentally releasing control and left and right hippocampal ablated pigeons from familiar training sites under anosmia, to render their navigational map dysfunctional, and after a phase-shift of the light-dark cycle, to place into conflict a pilotage-like landmark navigational strategy and a site-specific compass orientation landmark navigational strategy. Both left and right hippocampal ablated birds succeeded in learning to navigate by familiar landmarks, and both preferentially relied on sun-compass based, site-specific compass orientation to home. Like bilateral hippocampal lesioned birds, and in contrast to intact controls, neither ablation group adopted a pilotage-like strategy. We conclude that both the left and RHF are necessary if pilotage-like, familiar landmark navigation is to be learned or preferentially used for navigation.

Olfactory lateralization in homing pigeons: initial orientation of birds receiving a unilateral olfactory input

It has been shown that homing pigeons (Columba livia) rely on olfactory cues to navigate from unfamiliar locations. In fact, the integrity of the olfactory system, from the olfactory mucosa to the piriform cortex, is required for pigeons to navigate over unfamiliar areas. Recently it has been shown that there is a functional asymmetry in the piriform cortex, with the left piriform cortex more involved in the use of the olfactory navigational map than the right piriform cortex. To investigate further the lateralization of the olfactory system in relation to navigational processes in carrier pigeons, we compared their homing performance after either their left or the right nostril was plugged. Contrary to our expectations, we observed an impairment in the initial orientation of the pigeons with their right nostril plugged. However, both groups released with one nostril plugged tended to be poorer than control pigeons in their homing performance. The observed asymmetry in favour of the right nostril might be due to projections from the olfactory bulbs to the contralateral globus pallidum, a structure involved in motor responses.

Double egg deposition in the European pond turtle, Emys orbicularis, from central Italy.

Abstract The reproductive condition of Emys orbicularis females (presence or absence of oviductal eggs) was studied by palpation and radiography in a coastal population of central Italy (Tuscany). Forty‐nine females were recaptured at least twice during the reproductive period in 1997; 15 of them were found without eggs, 29 reproduced at least once, and 5 reproduced twice. The period of egg retention was estimated to be three to four weeks, and the time between two depositions about 34 days (n = 5). Despite the small sample of females reproducing twice, there is a significant tendency regarding a positive correlation between frequency of reproduction and female body size, within the study population. Reproductive females were bigger than females found without eggs; in addition, carapace length increased significantly in females found zero, one and two times with eggs. A comparison with preliminary data from northern populations of E. orbicularis, lead to the hypothesis that the annual clutch frequency cou...

Geographic variation of sexual size dimorphism and genetics in the European pond turtle, Emys orbicularis and Emys trinacris, of Italy

We studied the extent of sexual size dimorphism (SSD) in the European pond turtle throughout most of its Italian distribution range using eight morphological parameters of 580 Emys specimens, describing populations from Italy and large Mediterranean islands. Sexual size dimorphism was evident in the whole sample, but not in all the considered areas. SSD in studied Emys likely mirrors different growth trajectories and body size architectures between sexes at adult stage. Italian populations resulted separated into five groups with morphological differences: (i) East Po plain Italy north to Apennines; (ii) Tyrrhenian Italy; (iii) Corse; (iv) Sardinia; and (v) southern Italy and Sicily. These results agree with previously reported data on systematics of Mediterranean populations. Some Italian populations were also tested using restriction endonuclease analysis of 18+28S ribosomal DNA on 43 specimens of eight localities. The occurrence of a restriction fragment length polymorphism of rDNA was also noted. Morphometric and molecular data revealed that the two Calabrian populations were significantly differentiated with respect to the other sampled populations. According to previous mitocondrial DNA analyses, our results also support the view that the southern Italy populations of E. orbicularis are different from other Italian populations, even if our results do not perfectly fit with previous works.

Development of the navigational map in homing pigeons: effects of flight experience on orientation performance

Abstract Experiments have shown that pigeons, Columba livia , are able to develop navigational abilities even if reared and kept confined in an aviary, provided that they are exposed to the natural winds. In contrast, pigeons reared in a wind-screened aviary do not learn to navigate. Pigeons kept confined in a screened aviary when young do not learn to navigate even if, once they become adult, the screens are removed and the birds are exposed to natural winds for a period sufficiently long for map learning. In all of these experiments, pigeons were kept confined until the day of test release. In the present study we found that pigeons reared in a wind-screened aviary developed navigational abilities if, once adult, they were allowed to perform spontaneous flights around the loft. Nevertheless, their navigational performance never reached the level of controls. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Effects of monocular viewing on orientation in an arena at the release site and homing performance in pigeons.

Orientation and homing performance of pigeons with the left or right eye occluded were assessed in an arena at the release site and during the subsequent homing flight. Three release sites near Pisa, Italy, were used. Compared to binocular controls, monocular birds showed a bias in orientation towards the side of the viewing eye. In the arena, this bias was considerable and the mean deviation corresponded to the angle of the optical axis, suggesting a systematic error in visual representation during directional orientation. During flight after leaving the arena the directional bias decreased and the homeward orientation increased. While there was a slight lateralization of overall homing performance in favour of the right eye, there was no lateralization in directional orientation in the arena or at vanishing. Our results show that navigational mechanisms in either brain hemisphere profit from information obtained before take off and while flying over the release site. The existence and degree of lateralization is discussed in comparison to other studies that investigated homing under monocular viewing conditions.

Do bearing magnets affect the extent of deflection in clock-shifted homing pigeons?

When released after clock-shift, homing pigeons fail to orient towards the home direction but display a consistent deflection of their initial orientation due to the difference between the real sun azimuth and the computed azimuth according to the subjective time of each single bird. It has been reported that the size of the observed deflection is frequently smaller than expected and a discussion on the possible factors affecting the size of deflection has emerged. Some authors have proposed that the major factor in reducing the deflection after clock-shift is the simultaneous use of both the magnetic and the sun compasses, giving true and erroneous information, respectively, about the home direction. Therefore, a magnetic disturbance, by impeding the use of the geomagnetic information in determining the home direction, is presumed to increase the size of the deflection up to the levels of the expectation. To test this hypothesis, we released three groups of clock-shifted birds from unfamiliar locations (unmanipulated pigeons, pigeons bearing magnets on their head, and pigeons bearing magnets on their back) together with a group of unshifted control birds. As no difference in the orientation of the three groups emerged, we were not able to confirm the hypothesis of the role of the magnetic compass in reducing the expected deflection after clock-shift.