Toepel Ulrike, Knebel Jean-François, Hudry Julie, le Coutre Johannes, Murray Micah M.
How the human brain recognizes and differentiates objects from one another and likewise groups them into categories according to common features/actions has been a topic of neuropsychological and neuroscientific investigation for several decades. For example, ventral posterior temporal cortices have been found to play a major role not only in object processing in general (see Tanaka, 1997; Ungerleider and Haxby, 1994 for reviews), but also in category-selective processing (Moore and Price, 1999; Peissig and Tarr, 2007; Perani et al., 1995, 1999). One axis along which objects appear to be differentiated is whether their referent is something living or an artifact (i.e., manufactured). This distinction appears to hold true for visually presented stimuli (e.g., Caramazza and Shelton, 1998; Gerlach, 2007; Martin, 2007) as well as their auditory counterparts (Lewis et al., 2005; Murray et al., 2006). In addition to this categorical distinction, there is abundant evidence that the processing of faces (and places) may recruit highly specialized neural circuitry (e.g., Bentin et al., 2007; Kanwisher and Yovel, 2006). One perspective is that such specialized responsiveness follows from the social importance of faces in human interactions. More recently, investigations have revealed that objects and words can also be differentially processed according to their associated actions by regions of the ventral premotor cortex and posterior parietal areas (see Culham and Valyear, 2006; Johnson-Frey, 2004; Lewis et al., 2005; Pulvermuller, 2005 for reviews). Applying a similar line of reasoning, other socially or biologically important object categories would also be predicted to engage specialized neural circuitry. Foods and their proper discrimination, for example, are of utmost importance for an organism’s survival. While this object category has been the subject of comparatively few functional neuroimaging studies, there is evidence to suggest that food images may activate brain regions distinct from those activated by other object categories (Killgore et al., 2003; LaBar et al., 2001; Rothemund et al., 2007; Santel et al., 2006; Simmons et al., 2005). In particular, viewing pictures of foods as opposed to nonfoods has been shown to activate frontal cortices as well as the frontal operculum and the insula (Killgore et al., 2003; LaBar et al., 2001; Simmons et al., 2005), the latter thought to be primary gustatory cortex regions (O’Doherty et al., 2001). Viewing foods can also modulate the activity in secondary gustatory regions within the orbitofrontal cortex (OFC; Beaver et al., 2006; Simmons et al., 2005). In addition to this, wider network of brain regions, striate, and extrastriate visual areas also exhibit modulated activity during the categorization of foods from tools (Killgore et al., 2003; LaBar et al., 2001; Simmons et al., 2005). However, from this evidence alone the basis upon which foods are discriminated from other objects and categorized among each other cannot be determined. For example, it may be either the reward and/or sensory features of the stimuli that drive the observed differential responses. Also, from these data it cannot be discerned which area(s) among the network of regions exhibiting differential responses is first involved and in what sequence other regions contribute to the discrimination of foods. Rather, temporal information is essential for constructing accurate models of food discrimination and evaluation. Electrical neuroimaging based on electroencephalographic recordings has been proven to be a powerful tool to identify the spatiotemporal dynamics during the categorization of other object categories than food (e.g., tools, animals, vehicles, and faces). It has been shown that categorization and in some cases within-category discrimination occurs within the initial 100–200 ms following stimulus presentation (Bentin et al., 2007; Eger et al., 2003; Fabre-Thorpe et al., 2001; Ji et al., 1998; Johnson and Olshausen, 2003; Kanwisher and Yovel, 2006; Michel et al., 2004a; Pizzagalli et al., 1999; Proverbio et al., 2007; Thorpe et al., 1996; VanRullen and Thorpe, 2001). Given this evidence, we hypothesized that the discrimination of food from nonfood images would occur within a similar time frame as that of other object categories. Moreover, although food is quintessential for survival, inherent differences exist with respect to nutritional and energetic value as well as hedonic attributes. As such, even though energetic value and palatability are critical factors influencing eating behavior, decisions regarding food intake are often guided by factors that can be detrimental for an individual’s health. Often, high-fat foods are consumed with more pleasure, and in larger quantities, than low-fat foods. One possibility is that the drive for hedonic experiences may lead to inappropriate food consumption behavior. In turn, overeating of detrimental foods can ultimately result in obesity, diabetes, and hypertension, which are increasingly prevalent among industrialized societies. Understanding the decision processes leading to food selection and consumption, and in particular how the brain appraises the presentation of foods before they are ingested, is likely to prove essential for learning how to control and correct inappropriate eating behavior.
几十年来,人类大脑如何识别和区分不同物体,并根据共同特征/动作将它们归类,一直是神经心理学和神经科学研究的课题。例如,已发现腹侧后颞叶皮层不仅在一般物体处理中起主要作用(见Tanaka,1997;Ungerleider和Haxby,1994的综述),而且在类别选择性处理中也起主要作用(Moore和Price,1999;Peissig和Tarr,2007;Perani等人,1995,1999)。物体区分的一个维度是其指代物是生物还是人工制品(即制造出来的)。这种区分对于视觉呈现的刺激(如Caramazza和Shelton,1998;Gerlach,2007;Martin,2007)以及它们的听觉对应物(Lewis等人,2005;Murray等人,2006)似乎都成立。除了这种类别区分外,有大量证据表明,面部(和地点)的处理可能会调用高度专门化的神经回路(如Bentin等人,2007;Kanwisher和Yovel,2006)。一种观点认为,这种专门的反应性源于面部在人际互动中的社会重要性。最近的研究表明,腹侧运动前皮层和顶叶后部区域也可以根据物体的相关动作对物体和单词进行不同的处理(见Culham和Valyear,2006;Johnson-Frey,2004;Lewis等人,2005;Pulvermuller,2005的综述)。运用类似的推理思路,可以预测其他具有社会或生物学重要性的物体类别也会调用专门的神经回路。例如,食物及其正确辨别对生物体的生存至关重要。虽然这个物体类别相对较少成为功能性神经成像研究的主题,但有证据表明食物图像可能会激活与其他物体类别激活区域不同的脑区(Killgore等人,2003;LaBar等人,2001;Rothemund等人,2007;Santel等人,2006;Simmons等人,2005)。特别是,与非食物相比,观看食物图片已被证明会激活额叶皮层以及额下回和脑岛(Killgore等人,2003;LaBar等人,2001;Simmons等人,2005),后者被认为是主要的味觉皮层区域(O’Doherty等人,2001)。观看食物还可以调节眶额皮层(OFC)内次级味觉区域的活动(Beaver等人,2006;Simmons等人,2005)。除此之外,在将食物与工具进行分类时,更广泛的脑区网络、纹状和纹外视觉区域也会表现出活动调制(Killgore等人,2003;LaBar等人,2001;Simmons等人,2005)。然而,仅从这些证据中无法确定食物与其他物体区分以及在彼此之间分类的基础。例如,驱动观察到的差异反应的可能是刺激的奖励和/或感官特征。同样,从这些数据中无法辨别在表现出差异反应的区域网络中,哪些区域首先参与,以及其他区域以何种顺序对食物辨别做出贡献。相反,时间信息对于构建准确的食物辨别和评估模型至关重要。基于脑电图记录的电神经成像已被证明是识别除食物之外其他物体类别(如工具、动物、车辆和面部)分类过程中时空动态的有力工具。研究表明,分类以及在某些情况下类别内的辨别在刺激呈现后的最初100 - 200毫秒内发生(Bentin等人,2007;Eger等人,2003;Fabre-Thorpe等人,2001;Ji等人,1998;Johnson和Olshausen,2003;Kanwisher和Yovel,2006;Michel等人,2004a;Pizzagalli等人,1999;Proverbio等人,2007;Thorpe等人,1996;VanRullen和Thorpe,2001)。基于此证据,我们假设食物与非食物图像的辨别将在与其他物体类别相似的时间范围内发生。此外,尽管食物对生存至关重要,但在营养和能量价值以及享乐属性方面存在内在差异。因此,尽管能量价值和适口性是影响饮食行为的关键因素,但关于食物摄入的决定通常受可能对个体健康有害的因素指导。通常,高脂肪食物比低脂肪食物更令人愉悦地被食用,且食用量更大。一种可能性是对享乐体验的追求可能导致不适当的食物消费行为。反过来,有害食物的过度食用最终可能导致肥胖、糖尿病和高血压,这些疾病在工业化社会中越来越普遍。理解导致食物选择和消费的决策过程,特别是大脑在食物摄入前如何评估食物的呈现,对于学习如何控制和纠正不适当的饮食行为可能至关重要。