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      Use of Unpalatable Forages by Ruminants: The Influence of Experience with the Biophysical and Social Environment

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          Abstract

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          Unpalatable forages, due to either low nutrient content or the presence of toxic compounds, are widespread and represent a challenge for ruminant nutrition, health, and welfare. If we find ways to encourage consumption of unpalatable forages, they could provide at least part of the nutrient requirements of ruminants. Our objective was to synthesize the role of diverse environmental experiences on the use of unpalatable forages by ruminants. Experimental evidence shows that experience can alter both morpho-physiological and psychological (learning) mechanisms to better cope with unpalatable forages, particularly early in life when body functions are more amenable to change. Furthermore, experiential learning provides flexibility in diet selection, which is critical in changing foraging environments. By understanding and applying behavioural principles, it is possible to better devise management plans that optimize the nutrition, health, and welfare of herbivores grazing unpalatable forages throughout their life. In addition, a more uniform use of resources can be achieved from the landscape level down to the individual plant, with consequent benefits to ecosystem integrity and stability.

          Abstract

          Unpalatable forage resources (low nutrient density, potentially toxic metabolites) are widespread and represent a challenge for ruminant nutrition, health, and welfare. Our objective was to synthesize the role of biophysical and social experience on the use of unpalatable forages by ruminants, and highlight derived behavioural solutions for the well-being of soils, plants, and animals. Environmental experiences early in life modulate gene expression and promote learning, which alters morpho-physiological and psychological mechanisms that modify behavioural responses and change food and habitat selection. In this process, ruminants can become better adapted to the habitat where they are reared. Moreover, experiential learning provides flexibility in diet selection, which is critical for changing foraging environments. Learned associations between unpalatable and palatable foods, if ingested in appropriate amounts, sequence, and close temporal association, induce the development of preference for the former type of food. In this way, a more uniform use of resources can be achieved from the landscape level down to the individual plant, with the associated benefits to ecosystem integrity and stability. Ruminants can also learn the medicinal benefits of ingesting foods with toxins (e.g., condensed tannins and saponins with antiparasitic properties). This knowledge on behavioural processes can be translated into behavioural applications that provide low-cost solutions to many challenges that producers face in managing sustainable livestock production systems.

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          Epigenetics for ecologists.

          There is now mounting evidence that heritable variation in ecologically relevant traits can be generated through a suite of epigenetic mechanisms, even in the absence of genetic variation. Moreover, recent studies indicate that epigenetic variation in natural populations can be independent from genetic variation, and that in some cases environmentally induced epigenetic changes may be inherited by future generations. These novel findings are potentially highly relevant to ecologists because they could significantly improve our understanding of the mechanisms underlying natural phenotypic variation and the responses of organisms to environmental change. To understand the full significance of epigenetic processes, however, it is imperative to study them in an ecological context. Ecologists should therefore start using a combination of experimental approaches borrowed from ecological genetics, novel techniques to analyse and manipulate epigenetic variation, and genomic tools, to investigate the extent and structure of epigenetic variation within and among natural populations, as well as the interrelations between epigenetic variation, phenotypic variation and ecological interactions.
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            Effects of the Social Environment and Stress on Glucocorticoid Receptor Gene Methylation: A Systematic Review.

            The early-life social environment can induce stable changes that influence neurodevelopment and mental health. Research focused on early-life adversity revealed that early-life experiences have a persistent impact on gene expression and behavior through epigenetic mechanisms. The hypothalamus-pituitary-adrenal axis is sensitive to changes in the early-life environment that associate with DNA methylation of a neuron-specific exon 17 promoter of the glucocorticoid receptor (GR) (Nr3c1). Since initial findings were published in 2004, numerous reports have investigated GR gene methylation in relationship to early-life experience, parental stress, and psychopathology. We conducted a systematic review of this growing literature, which identified 40 articles (13 animal and 27 human studies) published since 2004. The majority of these examined the GR exon variant 1F in humans or the GR17 in rats, and 89% of human studies and 70% of animal studies of early-life adversity reported increased methylation at this exon variant. All the studies investigating exon 1F/17 methylation in conditions of parental stress (one animal study and seven human studies) also reported increased methylation. Studies examining psychosocial stress and psychopathology had less consistent results, with 67% of animal studies reporting increased exon 17 methylation and 17% of human studies reporting increased exon 1F methylation. We found great consistency among studies investigating early-life adversity and the effect of parental stress, even if the precise phenotype and measures of social environment adversity varied among studies. These results are encouraging and warrant further investigation to better understand correlates and characteristics of these associations.
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              Social Learning in Animals: Empirical Studies and Theoretical Models

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                Author and article information

                Journal
                Animals (Basel)
                Animals (Basel)
                animals
                Animals : an Open Access Journal from MDPI
                MDPI
                2076-2615
                14 April 2018
                April 2018
                : 8
                : 4
                : 56
                Affiliations
                [1 ]Centro de Recursos Naturales Renovables de la Zona Semiárida (CERZOS), Departamento de Agronomía, Universidad Nacional del Sur (UNS)-CONICET, La carrindanga km 7, Bahía Blanca 8000, Argentina
                [2 ]Department of Wildland Resources, Quinney College of Natural Resources, Utah State University, Logan, UT 84322-5230, USA; juan.villalba@ 123456usu.edu
                Author notes
                [* ]Correspondence: cedistel@ 123456criba.edu.ar ; Tel.: +54-291-486-1124
                Article
                animals-08-00056
                10.3390/ani8040056
                5946140
                29662017
                be99bd26-4897-473a-a649-a126e0782d60
                © 2018 by the authors.

                Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( http://creativecommons.org/licenses/by/4.0/).

                History
                : 09 March 2018
                : 11 April 2018
                Categories
                Review

                unpalatable forages,ruminants,environmental experience,early experience,diet selection,habitat selection

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