理解进化中的符号过程-CSDN博客

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注：机翻，未校。

Comprehending the Semiosis of Evolution

Published: 04 April 2016

Volume 9, pages 1–6, (2016)

Alexei Sharov, Timo Maran & Morten Tønnessen

Most contemporary evolutionary biologists consider perception, cognition, and communication just like any other adaptation to the environmental selection pressures. A biosemiotic approach adds an unexpected turn to this Neo-Darwinian logic and focuses not so much on the evolution of semiosis as it does on the semiosis of evolution. What is meant here, is that evolutionary forces are themselves semiotically constrained and contextualized. The effect of environmental conditions is always mediated by the responses of organisms, who select their developmental pathways and actions based on heritable or memorized past experience and a variety of external and internal signals. In particular, recognition and categorization of objects, learning, and communication (both intraspecific and interspecific) can change the evolutionary fate of lineages. Semiotic selection, an effect of choice upon other species (Maran and Kleisner 2010), active habitat preference (Lindholm 2015), making use of and reinterpreting earlier semiotic structures – known as semiotic co-option (Kleisner 2015), and semiotic scaffolding (Hoffmeyer 2015; Kull 2015), are some further means by which semiosis makes evolution happen.
大多数当代进化生物学家认为感知、认知和交流就像任何其他对环境选择压力的适应一样。生物符号学方法为这种新达尔文主义逻辑增加了一个意想不到的转折，它与其说关注符号学的进化，不如说关注进化的符号学。这里的意思是，进化力量本身在符号学上受到限制和语境化。环境条件的影响总是由生物体的反应介导的，生物体根据可遗传或记忆的过去经验以及各种外部和内部信号来选择它们的发育途径和行动。特别是，物体的识别和分类、学习和交流（种内和种间）可以改变谱系的进化命运。符号学选择，选择对其他物种的影响（Maran 和 Kleisner 2010），活跃的栖息地偏好（Lindholm 2015），利用和重新解释早期的符号学结构——称为符号学共选（Kleisner 2015）和符号学支架（Hoffmeyer 2015;Kull 2015），是符号学使进化发生的一些进一步手段。

Semiotic processes are easily recognized in animals that communicate and learn, but it is difficult to find directly analogous processes in organisms without nerves and brains. Molecular biologists are used to talk about information transfer via cell-to-cell communication, DNA replication, RNA or protein synthesis, and signal transduction cascades within cells. However, these informational processes are difficult to compare with perception-related sign processes in animals because information requires interpretation by some agency, and it is not clear where the agency in cells is. In bacterial cells, all molecular processes appear deterministic, with every signal, such as the presence of a nutrient or toxin, launching a pre-defined cascade of responses targeted at confronting new conditions. These processes lack an element of learning during the bacterial life span, and thus cannot be compared directly with complex animal and human semiosis, where individual learning plays a decisive role.
符号学过程在会交流和学习的动物中很容易识别，但在没有神经和大脑的生物体中很难找到直接类似的过程。分子生物学家用于讨论通过细胞间通讯、DNA 复制、RNA 或蛋白质合成以及细胞内的信号转导级联反应进行的信息传递。然而，这些信息过程很难与动物的感知相关标志过程进行比较，因为信息需要一些机构进行解释，并且不清楚细胞中的机构在哪里。在细菌细胞中，所有分子过程都显得具有确定性，每个信号（例如营养物质或毒素的存在）都会启动针对新条件的预定义级联反应。这些过程在细菌生命周期中缺乏学习元素，因此无法直接与复杂的动物和人类符号学相提并论，其中个体学习起着决定性的作用。

The determinism of the molecular clockwork was summarized in the dogma that genes determine the phenotype and not the other way around. As a result, the Modern Synthesis (MS) theory presented evolution as a mechanical process that starts with blind random variation of the genome, and ends with automatic selection of the fittest phenotypes. Although this theory may explain quantitative changes in already existing features, it certainly cannot describe the emergence of new organs or signaling pathways. The main deficiency of such explanations is that the exact correspondence between genotypes and phenotypes is postulated a priori. In other words, MS was built like Euclidean geometry, where questioning the foundational axioms will make the whole system fall, like a house of cards.
分子发条的决定论总结为基因决定表型的教条，而不是相反。因此，现代合成（MS）理论将进化描述为一个机械过程，从基因组的盲目随机变异开始，到自动选择最适者表型结束。尽管该理论可以解释现有特征的定量变化，但它肯定不能描述新器官或信号通路的出现。这种解释的主要缺陷是基因型和表型之间的确切对应关系是先验假设的。换句话说，MS 的构建就像欧几里得几何一样，质疑基本公理将使整个系统倒塌，就像纸牌屋一样。

The discipline of biosemiotics has generated a new platform for explaining biological evolution. It considers that evolution is semiosis, a process of continuous interpretation and re-interpretation of hereditary signs alongside other signs that originate in the environment or the body. According to Hoffmeyer and Emmeche (1991: 144),
生物符号学学科为解释生物进化创造了一个新的平台。它认为进化是符号学，是对遗传符号以及源自环境或身体的其他符号的不断解释和重新解释的过程。根据 Hoffmeyer 和 Emmeche （1991： 144）的说法，

[e]xpressed in the metaphorics of language, the zygote ‘reads’ the ‘book’ in its DNA, ‘interprets’ its meaning as a kind of ‘manual’ for the construction of a tool for survival, the individual organism. With the help from this tool, the egg cell can hope to continue its cell-line for yet another generation on the condition, of course, that the tool is sufficiently well-made to survive and reproduce in its ecological niche. Implied in this view is a very important but widely overlooked fact: The DNA does not specify the zygote, the zygote must be there beforehand.”
受精卵在语言的隐喻中，在其 DNA 中“阅读”“这本书”，将其含义“解释”为一种构建生存工具的“手册”，即单个有机体。在这个工具的帮助下，卵细胞可以希望将其细胞系延续到另一代，当然，前提是该工具足够精良，可以在其生态位中生存和繁殖。这种观点暗示了一个非常重要但被广泛忽视的事实：DNA 没有指定受精卵，受精卵必须事先存在。

This view of evolution differs radically from widely accepted ideas in biology and science in general; and more explanations are therefore necessary. The first and main challenge is to define agency within living cells and demonstrate that this agency supports creative interpretation of signs. Molecular processes in cells are not deterministic but goal-directed. They are prone to errors, but most errors and faulty regulations are detected and become corrected or compensated in many different ways to minimize adverse effects on cellular functions, survival, and reproduction (Bruni 2008). As for learning capacity, this is not always seen within a single cell cycle; but learning certainly occurs in multi-generational lineages of reproducing cells, as exemplified by the, on our human timescale, rapid development of resistance to antibiotics in bacteria. Although multi-generational learning is supported by differential reproduction of genotypes, the evolution involved does not depend on a specific mutation happening. Any mutation out of hundreds could work equally well, if only it is interpreted properly in the context of cellular organization and helps to resolve the problem (Sharov 2014). Moreover, cells may initially utilize epigenetic memory to modify their functions, which may become enhanced later, genetically. Thus, living cells are agents capable of inventing new ways of living; in other words, they possess a degree of semiotic freedom (Hoffmeyer 2014).
这种进化论观点与生物学和一般科学中广泛接受的观点截然不同;因此，需要更多的解释。第一个也是主要的挑战是定义活细胞内的能动性，并证明这种能动性支持对符号的创造性解释。细胞中的分子过程不是确定性的，而是以目标为导向的。它们容易出错，但大多数错误和错误的规定都会被检测到，并以许多不同的方式得到纠正或补偿，以尽量减少对细胞功能、存活和繁殖的不利影响（Bruni 2008）。至于学习能力，这并不总是在单个细胞周期中看到;但学习肯定发生在繁殖细胞的多代谱系中，正如我们人类的时间尺度上细菌对抗生素耐药性的快速发展所证明的那样。尽管基因型的差异繁殖支持多代学习，但所涉及的进化并不取决于发生的特定突变。数百个突变中的任何突变都可以同样有效，只要它在细胞组织的背景下得到适当的解释并有助于解决问题（Sharov 2014）。此外，细胞最初可能利用表观遗传记忆来改变其功能，这些功能可能会在以后的遗传上得到增强。因此，活细胞是能够发明新生活方式的代理;换句话说，他们拥有一定程度的符号学自由度（Hoffmeyer 2014）。

The second problem in understanding the semiosis of evolution is the need to explain the emergence of interpretation capacity in primordial systems at the origin of life. Third, a scale of sophistication in semiotic systems has to be outlined, spanning from simple molecular signaling to creative cognition in humans and some animals. Fourth, we need to explain the remarkable consistency of phenotypic traits (i.e., heritability) in the absence of genetic determinism. The fifth challenge is to explain adaptability, which is the capacity to generate new solutions to problems of life in critical conditions. And the sixth problem is to describe the process of integration of living systems into higher-level super-systems or super-organisms. All these issues need to be addressed by biosemiotics, by convincingly identifying and describing the semiotic component at the different levels of biological organization. These six challenges are extensively discussed in this issue of Biosemiotics.
理解进化符号学的第二个问题是需要解释生命起源的原始系统中解释能力的出现。第三，必须概述符号学系统的复杂程度，从简单的分子信号传导到人类和一些动物的创造性认知。第四，我们需要解释在没有遗传决定论的情况下表型性状（即遗传力）的显着一致性。第五个挑战是解释适应性，即在危急条件下为生活问题产生新解决方案的能力。第六个问题是描述生命系统整合到更高层次的超级系统或超级有机体的过程。所有这些问题都需要通过生物符号学来解决，通过令人信服地识别和描述生物组织不同层次的符号学成分。这六大挑战在本期《生物符号学》中进行了广泛讨论。

The paper by Hoffmeyer and Stjernfelt elucidates steps in the evolution of semiotic competence in living organisms, which are: molecular recognition, prokaryote-eukaryote transformation, division of labor in multicellular organisms, emergence of phenotypic plasticity, sense perception, behavioral choice, active information gathering, collaboration, deception, learning and social intelligence, sentience, and consciousness. The initial step (molecular recognition) can be described in terms of physical and chemical relations, but yet qualifies for being called “semiotic” because it is embedded in the context of complex cellular processes, where physics and chemistry are only means for reaching the organism’s goals. This approach is based on the fundamental assumption of unity between life and semiosis: “life, semiosis, and agency make up one conceptual complex which, once realized in the wild, constitutes the basis of ongoing sophistication during evolution”. In contrast to semiosis, perception is viewed as a distinctively high-level activity that is constituted by a multitude of smaller-scale semiotic interactions. Sense perception is not limited to mental knowledge, but also includes bodily knowledge, which occurs even in brainless animals. The authors reject the idea that semiotic evolution started from iconic relations and then progressed into indexial and symbolic relations, because, in their view, all “semiotic processes typically include both iconic, indexical, and symbolic aspects”. They further argue that “even very simple sign processes [are always] truth related; [and] that the ability of informing an organism about aspects of environmental states-of-affairs, such as they truly are, forms the most basic raison-d’ê tre for signs in the first place.” In other words, a kind of propositional logic is claimed to be used by organisms from the very beginning of agency and life.
Hoffmeyer 和 Stjernfelt 的论文阐明了生物体符号学能力进化的步骤，这些步骤是：分子识别、原核生物-真核生物转化、多细胞生物中的分工、表型可塑性的出现、感觉感知、行为选择、主动信息收集、协作、欺骗、学习和社会智能、感知和意识。第一步（分子识别）可以用物理和化学关系来描述，但仍然有资格被称为“符号学”，因为它嵌入了复杂的细胞过程的背景下，其中物理和化学只是实现生物体目标的手段。这种方法基于生命和符号学之间统一的基本假设：“生命、符号学和能动性构成了一个概念复合体，一旦在野外实现，它就构成了进化过程中持续复杂的基础”。与符号学相反，感知被视为一种独特的高级活动，由大量较小规模的符号学交互构成。感官知觉不仅限于心理知识，还包括身体知识，这甚至发生在无脑的动物身上。作者拒绝了符号学进化从符号关系开始，然后发展到索引和符号关系的观点，因为在他们看来，所有“符号学过程通常包括符号、索引和符号方面”。他们进一步争辩说，“即使是非常简单的符号过程 [总是] 与真理相关;[并且] 告知生物体有关环境状况的各个方面的能力，例如它们的真实情况，首先构成了迹象最基本的理由。换句话说，一种命题逻辑被声称从能动性和生命的一开始就被有机体使用。

The paper by Giorgi and Bruni is focused on germ cells as minimal interpreters of hereditary signs that link subsequent generations of organisms into lineages. The authors summarize the properties and capacities of germ cells that give them semiotic agency. Namely, germ cells store a self-descriptive program and accumulate resources and molecular machinery that are necessary for interpretation of the genome and for self-construction. Biological evolution, then, is driven not just by changes in the genome, but also by changes in interpretation and in self-construction networks. These are not fully prescribed by the genome, and operate in a context-dependent way. In particular, germ cells modify their semiotic competence via interaction with the parental organism during cell migration and maturation. As a result, “germ cells come to constitute a channel of communication between the developing organism and the species[‘] genomic memory”. Development of embryos is regulated by communication between cells and organs, as well as by sensorial input from the environment. Structural and functional novelty can therefore emerge epigenetically.
Giorgi 和 Bruni 的论文专注于生殖细胞作为遗传迹象的最小解释者，这些遗传迹象将后代生物体连接到谱系中。作者总结了赋予它们符号学代理的生殖细胞的特性和能力。也就是说，生殖细胞存储一个自我描述的程序，并积累解释基因组和自我构建所必需的资源和分子机制。因此，生物进化不仅受到基因组变化的驱动，还受到解释和自我构建网络的变化的驱动。这些并非完全由基因组规定，并且以依赖于环境的方式运作。特别是，生殖细胞在细胞迁移和成熟过程中通过与亲本生物体的相互作用来改变其符号学能力。因此，“生殖细胞构成了发育中的生物体和物种[']基因组记忆之间的交流渠道”。胚胎的发育受细胞和器官之间的通讯以及来自环境的感觉输入的调节。因此，结构和功能新颖性可以通过表观遗传学出现。

Gilbert presents the process of embryo development as context-dependent interpretation of developmental signs. The context includes the previous history of the responding cell, and external factors. Cells therefore respond differently to the same signal (e.g. hormone or growth factor): some cells proliferate, some cells differentiate, and other cells die. In addition, organisms have evolved to alter their development in response to differences in temperature, diet, the presence of predators, or the presence of competitors. This plasticity of development allows organisms to generate a wide range of phenotypes on the basis of the same genotype. Some organisms have also evolved to expect developmental signals from symbionts, and these organisms develop abnormally if the symbiont signals are not present. These examples show that embryo development is a regulatory hub that supports both the heritability and the adaptability of the phenotype via integration of genetic, physiological, and ecological channels of communication.
吉尔伯特将胚胎发育过程描述为对发育迹象的上下文依赖性解释。上下文包括响应单元的先前历史记录和外部因素。因此，细胞对相同信号（例如激素或生长因子）的反应不同：一些细胞增殖，一些细胞分化，而另一些细胞死亡。此外，生物体已经进化到会根据温度、饮食、捕食者的存在或竞争对手的存在来改变它们的发育。这种发育的可塑性使生物体能够在相同基因型的基础上产生广泛的表型。一些生物体也已经进化到期待来自共生体的发育信号，如果共生体信号不存在，这些生物体就会异常发育。这些例子表明，胚胎发育是一个调节中心，通过整合遗传、生理和生态交流渠道来支持表型的遗传性和适应性。

The papers by Kull and Markoš respectively discuss the semiotic nature of biological species from different points of view. Kull argues that a species is a self-defining entity integrated by the capacity of organisms to recognize each other as potential mating partners. The logical category of species is based on “family resemblance”, as defined by Wittgenstein (membership in a category due to overlapping similarities), rather than on shared fulfillment of a common criterion. In contrast to Mayr, who viewed interbreeding as a criterion of species, Kull views mate recognition as a semiotic process that holds the species together in practice and in actual fact. Further, Kull develops a model where the extent of heritable variation in a population is controlled by the width of the recognition window that is accepted by individuals. Organisms mate successfully only if the difference in their features does not exceed the width of the recognition window. Highly diverse populations either reduce their variation in evolution via low mating capacity of strongly deviating individuals, or they segregate into multiple phenotypically distinct groups with preferential inter-group mating. This model explains sympatric speciation without inferring additional factors such as separation of niches. It also explains character displacement in the areas of species coexistence.
Kull 和 Markoš 的论文分别从不同的角度讨论了生物物种的符号学性质。Kull 认为，物种是一个自我定义的实体，由生物体相互识别为潜在交配伙伴的能力整合而成。物种的逻辑类别基于维特根斯坦定义的“家庭相似性”（由于重叠的相似性而属于某个类别），而不是基于共同标准的共同满足。与将杂交视为物种标准的 Mayr 相反，Kull 将配偶识别视为一种符号学过程，将物种在实践中和实际中结合在一起。此外，Kull 开发了一个模型，其中种群中可遗传变异的程度由个体接受的识别窗口的宽度控制。只有当生物体的特征差异不超过识别窗口的宽度时，它们才能成功交配。高度多样化的种群要么通过强烈偏离个体的低交配能力来减少它们在进化中的变异，要么它们分裂成多个表型不同的群体，优先进行群体间交配。该模型解释了同域物种形成，但没有推断其他因素，例如生态位的分离。它还解释了物种共存领域的特征位移。

Markoš develops an idea of Rappaport and Flegr, namely that the long-term evolution of biological species is isomorphic to the historical change of human cultures. Both processes are deeply rooted in the history, which is materialized in memory, experience, and internal dynamics. Both evolutionary and cultural changes result from continuous reinterpretation of conservative digital texts as well as from changes in the interpretative process itself. This semiotic process results in a continuous inventing of new ways of living.
马尔科什发展了拉帕波特和弗莱格尔的观点，即生物物种的长期进化与人类文化的历史变化是同构的。这两个过程都深深植根于历史中，而历史则体现在记忆、经验和内部动力中。进化和文化变化都是对保守数字文本的不断重新解释以及解释过程本身变化的结果。这个符号学过程导致了新生活方式的不断发明。

The paper by Turner is focused on the evolution of super-organisms, taking the termite colony as an example. The termite mound is a complex structure which has many important functions for the life of the colony. It is the surrounding environment for termite workers who inhabit, interpret, and rebuild it, and is the hereditary legacy of past generations of workers. Because abandoned mounds can be re-colonized, their structure provides an additional channel of long-term heredity. Turner describes the termite nest as a system with “swarm cognition”, where individual workers respond to chemical signs such as CO2 perturbations and recruit other workers for rebuilding the nest if necessary. The process of mound repair is regulated by feedback signals from the circulating air in the nest. Mound repair reveals a complex decision-making system at work in the termite colony. A mound, with its higher-level adaptations, can be understood as the body of a super-organism, and its repair process resembles wound healing.
特纳的论文侧重于超级生物的进化，以白蚁群为例。白蚁丘是一个复杂的结构，在白蚁群的生命周期中具有许多重要功能。它是白蚁工人居住、解释和重建它的周围环境，是过去几代工人的世袭遗产。因为废弃的土墩可以重新定殖，所以它们的结构提供了额外的长期遗传渠道。特纳将白蚁巢描述为一个具有“群体认知”的系统，其中单个工蚁对二氧化碳扰动等化学信号做出反应，并在必要时招募其他工蚁重建巢穴。土堆修复的过程由巢中循环空气的反馈信号调节。土墩修复揭示了白蚁群落中一个复杂的决策系统。土丘具有更高层次的适应性，可以理解为超级有机体的身体，其修复过程类似于伤口愈合。

Sharov describes biological evolution as preservation, advance, and emergence of functional information in natural agents (organisms, cells, molecular agents, populations, and symbiotic consortia). He defines functional information as a network of signs (including memory, internal messengers, and external signs) that are used by agents to preserve and regulate their functions. Organisms preserve functional information via active processes of copyingFootnote 1 and construction: the digital components are copied, whereas interpreting subagents, scaffolds, tools, and resources are constructed. The advance of functional information includes improvement and combinatorial modification of already existing functions. Selective reproduction of agents at any level of a hierarchy helps to improve functions over time in varying environments. Under stress, agents can produce adaptive phenotypes very fast by utilizing complex regulatory pathways, which have been developed in a long-term evolution that intermittently included crisis events. Each new feature can then be tested in the context of a different body part or stage in a life cycle. Finally, the emergence of new functions is based on the reinterpretation of already existing information. These include cases of preadaptation/exaptation and the Baldwin effect. The major steps in the progressive evolution of functional information were protosemiosis, where signs correspond directly to actions, and eusemiosis, where agents associate signs with objects. Sharov assumes that primitive organisms bear no internal representations of objects; this capacity, he holds, emerged only with the appearance of eusemiosis. The notion of protosemiosis helps to explain the origin of life, because eusemiotic organisms are too complex to emerge spontaneously from non-living matter.
Sharov 将生物进化描述为自然代理（生物体、细胞、分子代理、种群和共生联盟）中功能信息的保存、推进和出现。他将功能信息定义为代理用来保存和调节其功能的符号网络（包括记忆、内部信使和外部符号）。生物体通过主动的复制过程来保存功能信息脚注 1 和构建：复制数字组件，同时构建解释子代理、脚手架、工具和资源。功能信息的进步包括对现有功能的改进和组合修改。在层次结构的任何级别选择性复制代理有助于在不同环境中随着时间的推移改进功能。在压力下，代理可以通过利用复杂的调节途径非常快速地产生适应性表型，这些途径是在间歇性包括危机事件的长期演变中发展起来的。然后，可以在生命周期中不同身体部位或阶段的上下文中测试每个新功能。最后，新功能的出现是基于对现有信息的重新解释。这些包括预适应/驱逐和 Baldwin 效应的情况。功能信息逐步进化的主要步骤是原型符号，其中符号直接与动作对应，以及优符号学，其中代理将符号与物体联系起来。沙罗夫假设原始生物没有物体的内部表征;他认为，这种能力只是随着 Eusemiosis的出现而出现的。原生符号学的概念有助于解释生命的起源，因为优释学生物体太复杂了，无法从非生命物质中自发出现。

The paper by Deacon shifts the discussion on evolution from traditional terms, such as replication, mutation, and selective retention of genes, to the more basic living functions. These include the generation and reproduction of organic forms via self-organizing and self-assembling molecular and cellular processes. These processes, in turn, require specific constraints and boundary conditions that need to be produced reciprocally by other self-organizing processes. Deacon describes how two or more self-organizing processes can be coupled so that they generate each other’s supportive boundary conditions. This coupling is a higher-order constraint, which constitutes a sign vehicle that is “interpreted” when it helps to make the form of a new physical system equipped with the same future competence. This semiotic-dynamical relation constitutes Darwin’s “several powers” that make evolution possible.
Deacon 的论文将关于进化的讨论从传统术语（如复制、突变和基因的选择性保留）转移到更基本的生命功能。这些包括通过自组织和自组装分子和细胞过程产生和复制有机形式。反过来，这些过程需要特定的约束和边界条件，这些约束和边界条件需要由其他自组织过程相互产生。Deacon 描述了两个或多个自组织过程如何耦合，以便它们产生彼此的支持边界条件。这种耦合是一个高阶约束，它构成了一个符号载体，当它有助于使一个新的物理系统的形式配备相同的未来能力时，它就会被 “解释”。这种符号学-动力学关系构成了达尔文的“几种力量”，使进化成为可能。

In recent years, the attention paid to evolution in biosemiotics has increased considerably (see e.g. Barbieri 2008; Markoš et al. 2009; Deacon et al. 2012). Comprehending the relationships between evolution, signs and life is essential for developing the theoretical foundations of the biosemiotic paradigm. It also assists biosemiotics in finding its position among the biological sciences. The emergence of evo-devo theory, the extended synthesis, epigenetic studies and other like-minded schools of thought in biology has produced fertile soil for future discussions. Our understanding evolves.
近年来，人们对生物符号学进化的关注大大增加（参见 Barbieri 2008;Markoš 等人，2009 年;Deacon 等人，2012 年）。理解进化、符号和生命之间的关系对于发展生物符号学范式的理论基础至关重要。它还帮助生物符号学在生物科学中找到自己的位置。进化论的出现、扩展综合、表观遗传学研究和其他志同道合的生物学思想流派为未来的讨论提供了肥沃的土壤。我们的理解在不断发展。

Notes

Specific features of copying are recursion and digital-type stability at the level of components (e.g., DNA monomers).

References

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Author information

Authors and Affiliations

Laboratory of Genetics and Genomics, National Institute on Aging, Baltimore, MD, USA

Alexei Sharov
Department of Semiotics, University of Tartu, Tartu, Estonia

Timo Maran
Department of Social Studies and Department of Health Studies, University of Stavanger, Stavanger, Norway

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