Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Effects of string length on the organization of rat string-pulling behavior


The string-pulling paradigm has been adapted to investigate many psychological phenomena across a range of animal species. Although varying string length has been shown to influence performance, the nature of the representation remains to be determined. Across three experiments, rats were shaped to pull string to receive food reinforcement. Either string length or reinforcement rate was manipulated to examine the influence on string-pulling behavior. Experiment 1 demonstrated that varied string length was sufficient to elicit an odor discrimination. Subsequent experiments provided evidence that varying string length (Experiment 2) and reinforcement rate (Experiment 3) produced qualitatively distinct patterns of string-pulling behavior. In Experiment 2 rats that received a long string were more likely to pull in the probe string to the end, yet no differences were observed in approach time between short and long groups. However, in Experiment 3 rats that received low reinforcement were less likely to pull in the probe string to the end and were slower to approach the string to begin pulling. These results are consistent with rats using temporal and motivational characteristics to guide responding during string-pulling behavior.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7


  1. Amsel A (1958) The role of frustrative nonreward in noncontinuous reward situations. Psychol Bull 55(2):102.

  2. Blackwell AA, Köppen JR, Whishaw IQ, Wallace DG (2018a) String-pulling for food by the rat: assessment of movement, topography and kinematics of a bilaterally skilled forelimb act. Learn Motiv 61:63–73.

  3. Blackwell AA, Widick WL, Cheatwood JL, Whishaw IQ, Wallace DG (2018b) Unilateral forelimb sensorimotor cortex devascularization disrupts the topographic and kinematic characteristics of hand movements while string-pulling for food in the rat. Behav Brain Res 338:88–100.

  4. Blankenship PA, Cherep LA, Donaldson TN, Brockman SN, Trainer AD, Yoder RM, Wallace DG (2015) Otolith dysfunction alter exploratory movement in mice. Behav Brain Res 325:1–11.

  5. Blankenship PA, Cheatwood JL, Wallace DG (2017) Unilateral lesions of the dorsocentral striatum (DCS) disrupt spatial and temporal characteristics of food protection behavior. Brain Struct Funct 222(6):2697–2710.

  6. Buhusi CV, Meck WH (2005) What makes us tick? Functional and neural mechanisms of interval timing. Nat Rev Neurosci 6(10):755.

  7. Capaldi EJ (1966) Partial reinforcement: a hypothesis of sequential effects. Psychol Rev 73(5):459–477.

  8. Cheng K, Spetch ML, Miceli P (1996) Temporal duration and spatial position. J Exp Psychol Anim Behav Process 22(2):175–182.

  9. Chung SH, Herrnstein RJ (1967) Choice and delay of reinforcement 1. J Exp Anal Behav 10(1):67–74.

  10. Crutchfield RS (1939) The determiners of energy expenditure in string-pulling by the rat. J Psychol 7(1):163–178.

  11. Davison M, McCarthy D (1988) The matching law: a research review. Erlbaum, Hillsdale, New Jersey

  12. Eilam D, Golani I (1989) Home base behavior of rats (Rattus norvegicus) exploring a novel environment. Behav Brain Res 34(3):199–211.

  13. Georgopoulos AP, Kalaska JF, Caminiti R, Massey JT (1982) On the relations between the direction of two-dimensional arm movements and cell discharge in primate motor cortex. J Neurosci 2(11):1527–1537.

  14. Golani I, Benjamini Y, Eilam D (1993) Stopping behavior: constraints on exploration in rats (Rattus norvegicus). Behav Brain Res 53(1–2):21–33.

  15. Herrnstein RJ (1961) Relative and absolute strength of response as a function of frequency of reinforcement 1, 2. J Exp Anal Behav 4(3):267–272.

  16. Jacobs IF, Osvath M (2015) The string-pulling paradigm in comparative psychology. J Comp Psychol 129(2):89.

  17. Kjelstrup KB, Solstad T, Brun VH, Hafting T, Leutgeb S, Witter MP et al (2008) Finite scale of spatial representation in the hippocampus. Science 321(5885):140–143.

  18. Lake JI, Meck WH (2013) Differential effects of amphetamine and haloperidol on temporal reproduction: dopaminergic regulation of attention and clock speed. Neuropsychologia 51(2):284–292.

  19. Martin MM, Winter SS, Cheatwood JL, Carter LA, Jones JL, Weathered SL et al (2008) Organization of food protection behavior is differentially influenced by 192 IgG-saporin lesions of either the medial septum or the nucleus basalis magnocellularis. Brain Res 1241:122–135.

  20. Matell MS, Berridge KC, Aldridge JW (2006) Dopamine D1 activation shortens the duration of phases in stereotyped grooming sequences. Behav Proc 71(2–3):241–249.

  21. McCoy DE, Schiestl M, Neilands P, Hassall R, Gray RD, Taylor AH (2019) New caledonian crows behave optimistically after using tools. Curr Biol 29(16):2737–2742.

  22. Meck WH (1996) Neuropharmacology of timing and time perception. Cogn Brain Res 3(3–4):227–242.

  23. Meck WH, Church RM (1987) Cholinergic modulation of the content of temporal memory. Behav Neurosci 101(4):457.

  24. Meck WH, Church RM, Wenk GL, Olton DS (1987) Nucleus basalis magnocellularis and medial septal area lesions differentially impair temporal memory. J Neurosci 7(11):3505–3511.

  25. Norman WD, McSweeney FK (1978) Matching, contrast, and equalizing in the concurrent lever-press responding of rats. J Exp Anal Behav 29(3):453–462.

  26. Roberts S (1981) Isolation of an internal clock. J Exp Psychol Anim Behav Process 7(3):242.

  27. Tchernichovski O, Golani I (1995) A phase plane representation of rat exploratory behavior. J Neurosci Methods 62(1–2):21–27.

  28. Wallace DG, Hamilton DA, Whishaw IQ (2006a) Movement characteristics support a role for dead reckoning in organizing exploratory behavior. Anim Cogn 9(3):219–228.

  29. Wallace DG, Wallace PS, Field E, Whishaw IQ (2006b) Pharmacological manipulations of food protection behavior in rats: evidence for dopaminergic contributions to time perception during a natural behavior. Brain Res 1112(1):213–221.

  30. Weinstock S (1954) Resistance to extinction of a running response following partial reinforcement under widely spaced trials. J Comp Physiol Psychol 47(4):318.

  31. Whishaw IQ, Gorny BP (1994) Food wrenching and dodging: eating time estimates influence dodge probability and amplitude. Aggress Behav 20(1):35–47.;2-S

  32. Whishaw IQ, Vanderwolf CH (1973) Hippocampal EEG, behavior: changes in amplitude and frequency of RSA (theta rhythm) associated with spontaneous and learned movement patterns in rats and cats. Behav Biol 8:461–484.

  33. Winter SS, Köppen JR, Ebert TBN, Wallace DG (2013) Limbic system structures differentially contribute to exploratory trip organization of the rat. Hippocampus 23(2):139–152.

Download references

Author information

Correspondence to Ashley A. Blackwell.

Additional information

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Blackwell, A.A., Wallace, D.G. Effects of string length on the organization of rat string-pulling behavior. Anim Cogn 23, 415–425 (2020).

Download citation


  • Time
  • Reinforcement rate
  • Odor
  • Discrimination
  • Probe trials
  • Movement kinematic