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Food quantity discrimination in puppies (Canis lupus familiaris)

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There is considerable evidence that animals are able to discriminate between quantities. Despite the fact that quantitative skills have been extensively studied in adult individuals, research on their development in early life is restricted to a limited number of species. We, therefore, investigated whether 2-month-old puppies could spontaneously discriminate between different quantities of food items. We used a simultaneous two-choice task in which puppies were presented with three numerical combinations of pieces of food (1 vs. 8, 1 vs. 6 and 1 vs. 4), and they were allowed to select only one option. The subjects chose the larger of the two quantities in the 1 vs. 8 and the 1 vs. 6 combinations but not in the 1 vs. 4 combination. Furthermore, the last quantity the puppies looked at before making their choice and the time spent looking at the larger/smaller amounts of food were predictive of the choices they made. Since adult dogs are capable of discriminating between more difficult numerical contrasts when tested with similar tasks, our findings suggest that the capacity to discriminate between quantities is already present at an early age, but that it is limited to very easy discriminations.
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Animal Cognition (2020) 23:703–710
https://doi.org/10.1007/s10071-020-01378-z
ORIGINAL PAPER
Food quantity discrimination inpuppies (Canis lupus familiaris)
MariaElenaMilettoPetrazzini1· FabioMantese2· EmanuelaPrato‑Previde3
Received: 16 November 2019 / Revised: 24 March 2020 / Accepted: 30 March 2020 / Published online: 6 April 2020
© Springer-Verlag GmbH Germany, part of Springer Nature 2020
Abstract
There is considerable evidence that animals are able to discriminate between quantities. Despite the fact that quantitative
skills have been extensively studied in adult individuals, research on their development in early life is restricted to a limited
number of species. We, therefore, investigated whether 2-month-old puppies could spontaneously discriminate between
different quantities of food items. We used a simultaneous two-choice task in which puppies were presented with three
numerical combinations of pieces of food (1 vs. 8, 1 vs. 6 and 1 vs. 4), and they were allowed to select only one option. The
subjects chose the larger of the two quantities in the 1 vs. 8 and the 1 vs. 6 combinations but not in the 1 vs. 4 combination.
Furthermore, the last quantity the puppies looked at before making their choice and the time spent looking at the larger/
smaller amounts of food were predictive of the choices they made. Since adult dogs are capable of discriminating between
more difficult numerical contrasts when tested with similar tasks, our findings suggest that the capacity to discriminate
between quantities is already present at an early age, but that it is limited to very easy discriminations.
Keywords Quantity discrimination· Dog· Puppies· Food choice task· Development
Introduction
Being able to correctly estimate whether one quantity is
larger or smaller than another confers advantages in sev-
eral ecological contexts and allows animals to cope with
problems in their natural environments. For instance, such a
competence is useful in regulating intergroup conflicts based
on the number of individuals in an opponent group, selecting
the best mating strategy according to the number of potential
mating partners, and reducing predation risk by joining a
larger group of social companions. Because of their high
adaptive value, it is not surprising that quantitative abilities
have been widely described both in vertebrate and in inverte-
brate species (Agrillo and Bisazza 2018; Geary etal. 2014).
The benefits associated with quantitative abilities are
probably most obvious in a foraging context in which
animals try to maximise the amount of energy which they
acquire (Stephens and Krebs 1986) by selecting larger food
sources. A wealth of studies has exploited the natural pref-
erence for larger amounts of food to investigate the quan-
titative abilities of several species. One commonly used
paradigm is the spontaneous choice task, in which animals
are offered two quantities of food items and are allowed to
choose only one option. Species as different as elephants
(Perdue etal. 2012), olive baboons (Barnard etal. 2013),
cats (Bánszegi etal. 2016), crows (Bogale etal. 2014), tor-
toises (Gazzola etal. 2018), and guppies (Lucon-Xiccato
etal. 2015) proved able to select sets that contained larger
quantities of food. However, their performances were ratio-
dependent, and their accuracy decreased as the ratio between
the quantities being compared increased (e.g., discriminating
between 3 vs. 12 items with a 0.25 ratio was easier than dis-
criminating between 3 vs. 5 items with a 0.6 ratio) (Ferrigno
and Cantlon 2017).
To date, longitudinal data on numerical competence are
only available in humans. There is clear evidence that our
species is endowed with a “numbers sense”, namely the abil-
ity to discriminate between sets of different number of items
(Dehaene 1997), which is supported by the “approximate
number system” (ANS) (Halberda and Feigenson 2008).
The ANS allows humans to discriminate between quantities
* Maria Elena Miletto Petrazzini
mariaelena.milettopetrazzini@gmail.com
1 School ofBiological andChemical Sciences, Queen Mary
University ofLondon, London, UK
2 Department ofAnimal andHuman Biology, University
ofTurin, Turin, Italy
3 Department ofPathophysiology andTransplantation,
University ofMilan, Milan, Italy
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... Since the chicks' response time and accuracy is ratio dependent when they process both large (e.g. 10 vs. 20) and small numbers (e.g. 2 vs. 3) it suggests they rely only on the AMS (Rugani et al. 2013(Rugani et al. , 2014. To our knowledge the only mammalian study on the ontogeny of quantity discrimination, apart from children, was done in puppies of the domestic dog (Canis lupus familiaris), where puppies' choices were found to be also ratio dependent, but where they discriminated ratios only below 0.25 (Miletto Petrazzini et al. 2020). In contrast, adult dogs spontaneously differentiate between a ratio of 0.6 (Miletto Petrazzini and Wynne 2016;Ward and Smuts 2007) and they can be trained to discriminate between objects with an area ratio of 0.8 (Rivas-Blanco et al. 2020). ...
... Eight of these were the same as those tested with adult cats (Bánszegi et al. 2016): 1 vs. 4 (ratio: 0.25), 1 vs. 3 (ratio: 0.33), 2 vs. 5 (ratio: 0.4), 1 vs. 2 (ratio: 0.5), 2 vs. 4 (ratio: 0.5), 3 vs. 5 (ratio: 0.6), 2 vs. 3 (ratio: 0.67), 3 vs. 4 (ratio: 0.75). Since a recent study on puppies reported the lack of differentiation between 1 vs. 4 pieces of food (Miletto Petrazzini et al. 2020), we added 1 vs. 6 (ratio: 0.17) and 1 vs. 9 (ratio: 0.11) to give the kittens a considerably easier tasks. We also added 2 vs. 6 (ratio: 0.33) and 4 vs. 6 (ratio: 0.67) comparisons to test whether the kittens would differentiate between larger numbers too. ...
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... In particular, the number of studies investigating dogs' quantitative abilities have been increasing in recent years 14 . Several studies have shown that dogs are capable of discriminating larger from smaller quantities and their ability is in agreement with the Weber's law [41][42][43][44][45][46][47] . Dogs are able to discriminate up to a ratio of 0.67 spontaneously 47 , and up to 0.80 if trained to do so 45 . ...
... It is worth noting that the time spent looking at the unconnected items before dogs' choice was positively correlated with their final choices. The positive correlation between looking behaviour and dogs' choices has been recently reported also by Miletto Petrazzini et al. 46 in another food choice task. The fact that dogs looked at the unconnected stimulus and then selected this array indirectly resembles a phenomenon known in cognitive science as "quiet eye", defined as the final fixation or tracking gaze at a task-relevant location immediately prior to movement 62 . ...
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... 51 Likewise, most insights into quantity discrimination in dogs rose from visual numerical discrimination 52 Wynne, 2016, 2017;Prato-Previde et al., 2008). These studies showed that dogs, 53 including puppies (Petrazzini et al., 2020), tend to select significantly more often the visually presented 54 large food quantity than the small food quantity in a choice task Wynne, 2016, 2017) 55 or the visually presented larger amount of non-food items on two magnetic boards (Macpherson and 56 Roberts, 2013), particularly for quanities of lower ratio (i.e. a large difference between the two quantities). 57 This result conforms with Weber's law (Fechner, 1860), where two comparable quantities (i.e. when 58 the small quantity divided by the large quantity produces a large value) are harder to discriminate 59 than two different quantities (i.e. when the small quantity divided by the large quantity produces a 60 small value). ...
Preprint
Full-text available
The extraordinary olfactory capabilities in detection and rescue dogs are well-known. However, the olfactory performance varies by breed and search environment (Jezierski et al., 2014), as well as by the quantity of training (Horowitz et al., 2013). While detection of an olfactory cue inherently demands a judgment regarding the presence or absence of a cue at a given location, olfactory discrimination requires an assessment of quantity, a task demanding more attention and, hence, decreasing reliability as an informational source (Horowitz et al., 2013). This study aims at gaining more clarity on detection and discrimination of olfactory cues in untrained dogs and in a variety of dog breeds. Using a two-alternative forced choice (2AFC) paradigm, we assessed olfactory detection scores by presenting a varied quantity of food reward under one or the other hidden cup, and discrimination scores by presenting two varied quantities of food reward under both hidden cups. We found relatively reliable detection performances across all breeds and limited discrimination abilities, modulated by breed. We discuss our findings in relation to the cognitive demands imposed by the tasks and the cephalic index of the dog breeds.
Chapter
Davila and colleagues cover methods and review findings regarding dog olfactory cognition. The chapter begins by introducing the topic of dog cognition. Specific aspects of dog olfactory cognition are explored. The authors review methods and procedures that investigate different aspects of olfaction and memory, including episodic memory, working memory, and long-term memory. Next, olfactory category and concept learning in dogs is reviewed followed by a discussion of the nature of olfactory representations in dogs, as well as studies that involve an olfactory search image. Finally, the last section reviews olfactory quantity judgments. The authors conclude by emphasizing the importance of dog cognition and suggesting a converging operation approach for future research.
Preprint
Full-text available
The manuscript is a review of an article about detection and discrimination abilities in dogs that are delineated by breed and also by cephalic index. The original authors used classic psychophysics (Weber's Just Noticeable Difference or JND), approximate number system (ANS), and Gestalt laws in psychology to measure and compare canine olfactory abilities. In the review, it is recommended that future studies use the Signal Detection Theory (SDT)'s concepts of noise and bias. Examples are used to illustrate recommendations. Additionally, collaboration with effective trainers of canine scent detection and discrimination dogs (as was done by Schoon in Holland) is recommended in order to better quantify and differentiate the type and quantities scents utilized to test dogs, as well as the circumstances under which the scents are presented.
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Quantitative aspects of the study of animal and human behaviour are increasingly relevant to test hypotheses and find empirical support for them. At the same time, photo and video cameras can store a large number of video recordings and are often used to monitor the subjects remotely. Researchers frequently face the need to code considerable quantities of video recordings with relatively flexible software, often constrained by species‐specific options or exact settings. BORIS is a free, open‐source and multiplatform standalone program that allows a user‐specific coding environment to be set for a computer‐based review of previously recorded videos or live observations. Being open to user‐specific settings, the program allows a project‐based ethogram to be defined that can then be shared with collaborators, or can be imported or modified. Projects created in BORIS can include a list of observations, and each observation may include one or two videos (e.g. simultaneous screening of visual stimuli and the subject being tested; recordings from different sides of an aquarium). Once the user has set an ethogram, including state or point events or both, coding can be performed using previously assigned keys on the computer keyboard. BORIS allows definition of an unlimited number of events (states/point events) and subjects. Once the coding process is completed, the program can extract a time‐budget or single or grouped observations automatically and present an at‐a‐glance summary of the main behavioural features. The observation data and time‐budget analysis can be exported in many common formats ( TSV , CSV , ODF , XLS , SQL and JSON ). The observed events can be plotted and exported in various graphic formats ( SVG , PNG , JPG , TIFF , EPS and PDF ).
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