Behavioral flexibility is related to exploration, but not boldness, persistence or motor diversity
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Abstract
Behavioral flexibility, the ability to change behavior when circumstances change based on learning from previous experience, is thought to play an important role in a species' ability to successfully adapt to new environments and expand its geographic range. However, behavioral flexibility is rarely directly tested at the individual level. This limits our ability to determine how it relates to other traits, such as exploration or persistence, that might also influence individual responses to novel circumstances. Without this information, we lack the power to predict which traits facilitate a species' ability to adapt behavior to new environments. We use great-tailed grackles (a bird species; hereafter “grackles”) as a model to investigate this question because they have rapidly expanded their range into North America over the past 140 years. We evaluated whether grackle behavioral flexibility (measured as color reversal learning) correlated with individual differences in the exploration of new environments and novel objects, boldness towards known and novel threats, as well as persistence and motor diversity in accessing a novel food source. We determined that exploration of a novel environment across two time points and persistence when interacting with several different novel apparatuses were repeatable in individual grackles. There was no relationship between exploration or persistence and the two components of flexibility - the rate of learning to prefer a color option in the reversal learning task, and the rate of deviating from a preferred option. However, grackles that underwent serial reversal training to experimentally increase behavioral flexibility were more exploratory in that they spent more time in close proximity to the novel environment relative to control individuals. This indicates that, the more an individual investigated a novel apparatus, the more it was able to potentially learn and update its knowledge of current reward contingencies to adapt behavior accordingly. Our findings improve our understanding of the traits that are linked with flexibility in a highly adaptable species. We highlight the importance of using multiple different methods for measuring boldness and exploration to evaluate consistency of performance and therefore the methodological validity. We also show a link between exploration and behavioral flexibility that could facilitate adaptation to novel environmental changes.
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Animal behavior, like other kinds of phenotypic plasticity, is crucial for survival and reproduction in environments that vary over space and time. Behaviors themselves may need to be flexible when the distributions of environmental conditions themselves change; for example, short-term weather patterns and long-term climate conditions are changing due to human activity, and behavioral flexibility will likely be key to some population persisting during these changes and others going extinct. Thus, measuring this flexibility is key to understanding which species may be resilient to climate change.
Measuring behavioral flexibility is tricky as different studies may define and measure it differently and yet other studies may measure similar kinds of flexibility yet call them different things. This so-called "jingle-jangle" issue suggests …
Animal behavior, like other kinds of phenotypic plasticity, is crucial for survival and reproduction in environments that vary over space and time. Behaviors themselves may need to be flexible when the distributions of environmental conditions themselves change; for example, short-term weather patterns and long-term climate conditions are changing due to human activity, and behavioral flexibility will likely be key to some population persisting during these changes and others going extinct. Thus, measuring this flexibility is key to understanding which species may be resilient to climate change.
Measuring behavioral flexibility is tricky as different studies may define and measure it differently and yet other studies may measure similar kinds of flexibility yet call them different things. This so-called "jingle-jangle" issue suggests that studies can more robustly measure a behavioral trait when they use multiple behavioral tests. An additional issue is that measuring behavioral traits that vary between individuals due to genetic or development effects, often referred to as "personality" traits, requires that those trait differences be repeatable across time and between individuals.
With these issues in mind, McCune et al. (2019) presented a preregistration for experiments using a population of great-tailed grackles to investigate how behavioral flexibility relates to other important traits that are known to vary across individuals including exploratory behavior, boldness, and persistence and motor diversity in accessing a new food source. Behavioral flexibility is measured by the rate of learning a color associated with reward after first learning an association with a different color. That preregistration was recommended by PCI Ecology (Van Cleve, 2019). Now, McCune et al. (2025) present results from this study and find that only exploration of a novel environment and persistence were statistically repeatable behaviors. Both behaviors were not significantly correlated with behavioral flexibility. However, grackles that were trained to perform better in the color reversal learning task were more exploratory. This association between behavioral flexibility and exploratory tendencies may have evolved in grackles to help them survive in new environments, which they have proven very capable of doing as they have expanded their range in North American over the last 140 years (Wehtje, 2003).
There are a few key features of McCune et al. (2025) to merit recommendation. First, the authors are intent on demonstrating careful behavioral research practices including, as evidenced above, preregistering their hypotheses and predictions and making available both the preregistered and final analysis code. Second, the study demonstrates a thorough attempt to address two aspects that bedevil behavioral research, namely the "jingle-jangle" issue and repeatability of traits across individuals. Even after measuring multiple features of boldness, exploratory, persistence, McCune et al. (2025) find that only a subset of the measured behaviors are repeatable and only a subset of those are associated with behavioral flexibility. This suggests that only thorough studies like McCune et al. (2025) can start to probe difficult to measure behavioral associations that may be key to understanding how species will respond to our changing world.
References
McCune K, Lukas D, MacPherson M, Logan CJ (2025) Behavioral flexibility is related to exploration, but not boldness, persistence or motor diversity. EcoEvoRxiv, ver.2 peer-reviewed and recommended by PCI Ecology https://doi.org/10.32942/X2H33F
Van Cleve, J. (2019) Probing behaviors correlated with behavioral flexibility. PCI Ecology, 100020. https://doi.org/10.24072/pci.ecology.100020
McCune K, Rowney C, Bergeron L, Logan CJ. (2019) Is behavioral flexibility linked with exploration, but not boldness, persistence, or motor diversity? (http://corinalogan.com/Preregistrations/g_exploration.html) In principle acceptance by PCI Ecology of the version on 27 Mar 2019
Wehtje, W. (2003) The range expansion of the great-tailed grackle (Quiscalus mexicanus Gmelin) in North America since 1880. Journal of Biogeography 30:1593–1607 https://doi.org/10.1046/j.1365-2699.2003.00970.x.
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