ReviewCardiac defense: From attention to action☆
Introduction
Organisms react physiologically to the presence of danger or threat in order to protect themselves from potential injury or death. Typical defense responses reported in the experimental and clinical literature include freezing, startle, fainting, and the fight–flight response. These various responses can be categorized into two general forms of defense (Lang et al., 2000): immobility and active defense. The protective function of both types of reactions is evident. However, if too intense or prolonged, this defensive reactivity may constitute a serious risk for both mental and physical health. For many researchers and clinicians, excessive physiological reactivity is the main mechanism linking defense to stress and illness (Dienstbier, 1989, Lovallo and Gerin, 2003, Turner, 1994).
Fear and anxiety are the typical emotional reactions to the presence of danger and threat and are, therefore, closely linked to the concept of defense. Recent advances in the neurophysiology of fear and anxiety come primarily from research with animals using defense reactions such as freezing, startle, and escape-attack behaviors (Davis, 1992, LeDoux, 2000, Blanchard and Blanchard, 1989, Fanselow, 1994). Paradigms frequently used to study the neural circuit of fear (fear conditioning, fear sensitization, and fear potentiation) always include nociceptive or intense stimulation, eliciting unconditioned defense reactions, in order to investigate the neural pathways underlying the fear response. Research with humans has also made frequent use of intense or aversive stimulation in order to study fear modulation of specific protective reflexes such as eye-blink startle or postural freezing (Lang et al., 2000, Azevedo et al., 2005, Ruiz-Padial and Vila, 2007).
Pathological fear and anxiety are also closely related to defense. Perhaps, the most dramatic evidence of the relevance of defense in the development of pathological fear and anxiety comes from clinical studies on post-traumatic stress disorder as a consequence of extreme, life-threatening, traumatic events (Schauer et al., 2005). However, in these cases, as in any natural setting where a real danger emerges, it is difficult to talk of the defense reaction as a single entity. Rather, a dynamic sequence or cascade of defense reactions, from heightened attention to response mobilization, seems to take place depending primarily on the type and severity of the danger, its spatial and temporal proximity, and the success or failure of the initial defense responses to cope with it (Fanselow, 1994, Lang et al., 1997aa; McNaughton and Corr, 2004).
In this paper, we will focus on a specific defense reaction which has a long tradition in psychophysiological research: cardiac defense. We will begin with a brief review of the two opposite traditional approaches to understand this autonomic response: the cognitive and the motivational. Then we will present the classic model of cardiac defense and its basic assumptions concerning differentiation from other cardiac reflexes, namely orienting and startle. A critical analysis of these assumptions will follow centered on evidence from a systematic research of the cardiac response to intense acoustic stimulation. It will be shown that this response is characterized by a complex pattern of heart rate changes with accelerative and decelerative components, with sympathetic and parasympathetic mediating mechanisms, and with cognitive and motivational significance. Finally, an integrative model will be presented which emphasizes the dynamic nature of this defense reaction and provides a new framework in which the two opposite traditional approaches can be reconciled.
Section snippets
Traditional approaches to cardiac defense
The historical antecedents of the defense concept are rooted in the work of Pavlov (1927) and Cannon (1929). By the end of the nineteenth and beginning of the twentieth century, Pavlov and other Russian reflexologists (see Konorski, 1967) used the term defense reflex to refer to protective physiological responses elicited by noxious stimulation, such as hand withdrawal to an electric shock, eye blink to a puff of air, or vomit to bad food. Some years later, Cannon used the same term to refer to
The classic model on cardiac defense
The most elaborate theoretical model on cardiac defense was developed within the cognitive tradition. Based on the seminal work of Sokolov (1963) and Lacey and Lacey, 1958, Graham and Clifton, 1966 proposed a classification of cardiac reflexes that had a great impact in the field of psychophysiology. The classification was later extended and refined by Graham, 1979, Graham, 1992; (see also Graham and Hackley 1991) who proposed four basic reflexes elicited by stimuli as a function of their
A systematic search on cardiac defense
In the following sections we will summarize a series of studies on the heart rate component of the defense response in humans using as basic paradigm the presentation of an unexpected intense noise under different stimulus and task conditions. Instructions typically inform participants that they would hear brief intense noises but no indication of the exact moment of their presentation is given. The main dependent variable was always the beat-by-beat heart rate response recorded from the
The defense cascade
The description of cardiac defense to intense acoustic and electrocutaneous stimulation as a response pattern with both accelerative and decelerative components, with both sympathetic and parasympathetic influences, and with both attentional and motivational significance highlights the dynamic character of the defense reaction, obvious in natural settings. As has been emphasized by animal researchers (Blanchard and Blanchard, 1989, Fanselow, 1994), in natural settings such as the imminence of a
Conclusion
The studies reported here shows that, contrary to the classic model, cardiac defense to intense acoustic stimulation is a complex pattern of heart rate changes with accelerative and decelerative components, with sympathetic and parasympathetic physiological mediation, and with cognitive and motivational significance. This response pattern is congruent with an interpretation of the defense reaction as a sequence of responses from heightened attention to active defense. This interpretation helps
Acknowledgements
The various studies reported in this paper were supported by grants from the Spanish Ministry of Science and Education (projects PR82/1933, PB93-1096, PB97-0841, BSO2001-3211, and SEJ2004-07956), and the Junta de Andalucía (research group HUM-388).
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The present article is an updated and extended version of a previous article published in 2003 in The Spanish Journal of Psychology by Vila et al. entitled Cardiac Defense: Attention or Emotion? (The Spanish Journal of Psychology, 6, 60-78).