Why the carrot is more effective than the stick: different dynamics of punishment memory and reward memory and its possible biological basis

Neurobiol Learn Mem. 2009 Oct;92(3):370-80. doi: 10.1016/j.nlm.2009.05.003. Epub 2009 May 10.


One of the most extensively debated topics in educational psychology is whether punishment or reward is more effective for producing short-term and long-term behavioral changes, and it has been proposed that the effect of punishment is less durable than the effect of reward. However, no conclusive evidence to support this proposal has been obtained in any animals. We recently found that punishment memory decayed much faster than reward memory in olfactory learning and visual pattern learning in crickets. We also found that neurotransmitters conveying punishment and reward signals differ in crickets: dopaminergic and octopaminergic neurons play critical roles in conveying punishment and reward signals, respectively. In this study, we investigated whether these features are general features of cricket learning or are specific to olfactory and visual pattern learning. We found that crickets have the capability of color learning and that their color learning has the same features. Based on our findings in crickets and those reported in other species of insects, we conclude that these two features are conserved in many forms of insect learning. In mammals, aminergic neurons are known to convey reward and punishment signals in learning of a variety of sensory stimuli. We propose that the faster decay of punishment memory than reward memory observed in insects and humans reflects different cellular and biochemical processes after activation of receptors for amines conveying punishment and reward signals. The possible adaptive significance of relatively limited durability of punishment memory is proposed.

Publication types

  • Research Support, Non-U.S. Gov't

MeSH terms

  • Animals
  • Association Learning / drug effects
  • Association Learning / physiology
  • Color*
  • Conditioning, Classical / drug effects
  • Conditioning, Classical / physiology*
  • Dopamine Antagonists / pharmacology
  • Gryllidae / physiology*
  • Learning / drug effects
  • Learning / physiology*
  • Male
  • Memory / drug effects
  • Memory / physiology*
  • Models, Animal
  • Neuropsychological Tests
  • Photic Stimulation
  • Punishment*
  • Receptors, Biogenic Amine / antagonists & inhibitors
  • Reward*
  • Serotonin Antagonists / pharmacology
  • Time Factors


  • Dopamine Antagonists
  • Receptors, Biogenic Amine
  • Serotonin Antagonists
  • norsynephrine receptor