Gilda Santos Mühlen

Bio: Gilda Santos Mühlen is an academic researcher from Universidade Federal de Rondônia. The author has contributed to research in topics: Domestication & Hypocotyl. The author has an hindex of 7, co-authored 7 publications receiving 174 citations. Previous affiliations of Gilda Santos Mühlen include Federal University of Amazonas.

Evolution under domestication: contrasting functional morphology of seedlings in domesticated cassava and its closest wild relatives

Abstract: Although cassava (Manihot esculenta ssp. esculenta) is asexually propagated, farmers incorporate plants from seedlings into planting stocks. These products of sex are exposed to selection, which in agricultural environments should favour rapid growth. To examine whether seedling morphology has evolved under domestication, we compared domesticated cassava, its wild progenitor (M. esculenta ssp. flabellifolia) and their sister species (M. pruinosa) under controlled conditions. Field observations complemented laboratory study. In both wild taxa, the hypocotyl did not elongate (hypogeal germination) and cotyledons remained enclosed in the testa. In domesticated cassava, the hypocotyl elongated (epigeal germination), and cotyledons emerged and became foliaceous. The difference in hypocotyl elongation was fixed, whereas cotyledon morphology varied with environmental conditions in M. pruinosa. Comparative analysis suggests that epigeal germination is primitive in Manihot, that the lineage including wild ancestors of cassava evolved hypogeal germination--which confers greater tolerance to risks in their savanna environment--and that with domestication, there was a reversion to epigeal germination and photosynthetic cotyledons, traits conferring high initial growth rates in agricultural habitats.

Conteúdo cianogênico em progênies de mandioca originadas do cruzamento de variedades mansas e bravas

Abstract: As variedades de mandioca (Manihot esculenta Crantz ssp esculenta) sao classificadas pela taxonomia popular em bravas e mansas. As bravas tem sabor amargo, contem alto teor de glicosideos cianogenicos (superior a 100 mg de equivalente HCN/kg de polpa fresca de raiz) e sao consumidas apos serem processadas na forma de farinha, fecula e outros produtos. As mansas ou doces nao tem sabor amargo, contem baixo teor de glicosideos cianogenicos, sao consumidas com ou sem qualquer processamento, principalmente por meio de preparados domesticos simples. Neste trabalho, foram feitos cruzamentos para verificar o perfil cianogenico das progenies segregantes visando utilizar variedades bravas no melhoramento de mandiocas mansas. Duas variedades mansas (IAC 289-70 e IAC 576-70) e uma brava (SRT-1330 Xingu) foram polinizadas com uma variedade mansa (SRT-797 Ouro do Vale). Dos cruzamentos entre variedades mansas, 85,7% dos genotipos apresentaram teores abaixo de 100 mg eq. HCN/kg de polpa fresca de raiz. Do cruzamento entre as variedades mansa e a brava, apenas 31,3% dos genotipos mostraram teores abaixo desse valor. A media das progenies foi muito proxima a media dos parentais. Em todos os cruzamentos, a maior frequencia de genotipos esteve na classe entre 50 e 100 mg eq. HCN/kg de polpa fresca de raiz, mesmo no cruzamento entre brava e mansa. Em todos os cruzamentos apareceram segregantes transgressivos para maior e menor potencial cianogenico em relacao a qualquer parental. O tipo de segregacao observada indica ser a cianogenese um carater com heranca quantitativa que, embora tenha algum grau de influencia ambiental, torna possivel a selecao de individuos com maior ou menor teor cianogenico devido a alta herdabilidade oriunda de ampla variabilidade genotipica. Assim, variedades bravas podem surgir como recombinantes de variedades mansas, sendo tambem possivel selecionar variedades com baixo teor cianogenico em cruzamentos de mandiocas mansas com mandiocas bravas.

The evolutionary fate of phenotypic plasticity and functional traits under domestication in manioc: changes in stem biomechanics and the appearance of stem brittleness

Abstract: Domestication can influence many functional traits in plants, from overall life-history and growth form to wood density and cell wall ultrastructure. Such changes can increase fitness of the domesticate in agricultural environments but may negatively affect survival in the wild. We studied effects of domestication on stem biomechanics in manioc by comparing domesticated and ancestral wild taxa from two different regions of greater Amazonia. We compared mechanical properties, tissue organisation and wood characteristics including microfibril angles in both wild and domesticated plants, each growing in two different habitats (forest or savannah) and varying in growth form (shrub or liana). Wild taxa grew as shrubs in open savannah but as lianas in overgrown and forested habitats. Growth form plasticity was retained in domesticated manioc. However, stems of the domesticate showed brittle failure. Wild plants differed in mechanical architecture between shrub and liana phenotypes, a difference that diminished between shrubs and lianas of the domesticate. Stems of wild plants were generally stiffer, failed at higher bending stresses and were less prone to brittle fracture compared with shrub and liana phenotypes of the domesticate. Biomechanical differences between stems of wild and domesticated plants were mainly due to changes in wood density and cellulose microfibril angle rather than changes in secondary growth or tissue geometry. Domestication did not significantly modify “large-scale” trait development or growth form plasticity, since both wild and domesticated manioc can develop as shrubs or lianas. However, “finer-scale” developmental traits crucial to mechanical stability and thus ecological success of the plant were significantly modified. This profoundly influenced the likelihood of brittle failure, particularly in long climbing stems, thereby also influencing the survival of the domesticate in natural situations vulnerable to mechanical perturbation. We discuss the different selective pressures that could explain evolutionary modifications of stem biomechanical properties under domestication in manioc.

Crop domestication in the upper Madeira River basin

Abstract: Most native Amazonian crops were domesticated in the periphery of the basin. The upper Madeira River basin is an important part of this periphery where several important crops were domesticated and others are suspected to have been domesticated or arrived early. Some of these crops have been reasonably well studied, such as manioc, peanut, peach palm, coca and tobacco, while others are not as well known, such as the hot peppers Capsicum baccatum and C. frutescens, and still others need confirmation, such as cocoyam and annatto. We review the information available for manioc, peach palm, Capsicum, peanut, annatto and cocoyam. The state-of-the-art for Capsicum frutescens, annatto and cocoyam is insufficient to conclude definitively that they were domesticated in the upper Madeira, while all the others have at least one of their origins or centers of diversity in the upper Madeira.

Genetic Diversity and Differentiation of Brazilian Bitter and Sweet Manioc Varieties (Manihot esculenta Crantz, Euphorbiaceae) Based on SSR Molecular Markers

Abstract: Manioc (Manihot esculenta Crantz) originated in Amazonia and is the main staple for more than 800 million people worldwide; it also had a fundamental role as a source of calories for many pre-Columbian peoples, especially in Amazonia, where it was domesticated. There are two major groups of manioc varieties: sweet varieties have low amounts of toxic substances (cyanogenic glycosides) and may be consumed with minimum processing, while bitter varieties have a high degree of toxicity and must be detoxified to be safe before consumption. These groups are outcomes of divergent selective pressures. Natural selection probably maintains large amounts of cyanogenic glycosides to serve as a plant defense when in cultivation. Human selection may reduce the toxicity of the plants when roots are directly consumed, but may be neutral when the roots are consumed after some kind of processing. Although farmers recognize the distinction of the two groups of varieties, the variation of cyanogenic glycosides is continuous among different varieties. Genetic differentiation between sweet and bitter varieties was detected with molecular markers, as well as different patterns of groupings of varieties from different regions of Brazil. The genetic distinctions suggest that the sweet varieties originated during the initial domestication in southwestern Amazonia and bitter varieties arose later during cultivation in Amazonia, as hypothesized by Arroyo-Kalin in a recent paper. They also suggest that these groups of varieties were dispersed independently, even though they are cultivated complementarily today, with sweet varieties in home-gardens and bitter varieties in swiddens.

Identification of a Functional Transposon Insertion in the Maize Domestication Gene tb1

Abstract: Genetic diversity created by transposable elements is an important source of functional variation upon which selection acts during evolution. Transposable elements are associated with adaptation to temperate climates in Drosophila, a SINE element is associated with the domestication of small dog breeds from the gray wolf and there is evidence that transposable elements were targets of selection during human evolution. Although the list of examples of transposable elements associated with host gene function continues to grow, proof that transposable elements are causative and not just correlated with functional variation is limited. Here we show that a transposable element (Hopscotch) inserted in a regulatory region of the maize domestication gene, teosinte branched1 (tb1), acts as an enhancer of gene expression and partially explains the increased apical dominance in maize compared to its progenitor, teosinte. Molecular dating indicates that the Hopscotch insertion predates maize domestication by at least 10,000 years, indicating that selection acted on standing variation rather than new mutation.

The evolutionary ecology of clonally propagated domesticated plants.

Abstract: Summary While seed-propagated crops have contributed many evolutionary insights, evolutionary biologists have often neglected clonally propagated crops. We argue that widespread notions about their evolution under domestication are oversimplified, and that they offer rich material for evolutionary studies. The diversity of their wild ancestors, the diverse ecologies of the crop populations themselves, and the intricate mix of selection pressures, acting not only on the parts harvested but also on the parts used by humans to make clonal propagules, result in complex and diverse evolutionary trajectories under domestication. We examine why farmers propagate some plants clonally, and discuss the evolutionary dynamics of sexual reproduction in clonal crops. We explore how their mixed clonal ⁄ sexual reproductive systems function, based on the sole example studied in detail, cassava (Manihot esculenta). Biotechnology is now expanding the number of clonal crops, continuing the 10 000-yr-old trend to increase crop yields by propagating elite genotypes. In an era of rapid global change, it is more important than ever to understand how the adaptive potential of clonal crops can be maintained. A key component of strategies for preserving this adaptive potential is the maintenance of mixed clonal/sexual systems, which can be achieved by encouraging and valuing farmer knowledge about the sexual reproductive biology of their clonal crops. Contents Summary 318 I. Domesticated plants as model systems in evolutionary biology: bringing clonally propagated crops into the fold 319

Convergent evolution and parallelism in plant domestication revealed by an expanding archaeological record

Abstract: Recent increases in archaeobotanical evidence offer insights into the processes of plant domestication and agricultural origins, which evolved in parallel in several world regions. Many different crop species underwent convergent evolution and acquired domestication syndrome traits. For a growing number of seed crop species, these traits can be quantified by proxy from archaeological evidence, providing measures of the rates of change during domestication. Among domestication traits, nonshattering cereal ears evolved more quickly in general than seed size. Nevertheless, most domestication traits show similarly slow rates of phenotypic change over several centuries to millennia, and these rates were similar across different regions of origin. Crops reproduced vegetatively, including tubers and many fruit trees, are less easily documented in terms of morphological domestication, but multiple lines of evidence outline some patterns in the development of vegecultural systems across the New World and Old World tropics. Pathways to plant domestication can also be compared in terms of the cultural and economic factors occurring at the start of the process. Whereas agricultural societies have tended to converge on higher population densities and sedentism, in some instances cultivation began among sedentary hunter-gatherers whereas more often it was initiated by mobile societies of hunter-gatherers or herder-gatherers.

Parallel Domestication of the Shattering1 Genes in Cereals

Abstract: A key step during crop domestication is the loss of seed shattering. Here, we show that seed shattering in sorghum is controlled by a single gene, Shattering1 (Sh1), which encodes a YABBY transcription factor. Domesticated sorghums harbor three different mutations at the Sh1 locus. Variants at regulatory sites in the promoter and intronic regions lead to a low level of expression, a 2.2-kb deletion causes a truncated transcript that lacks exons 2 and 3, and a GT-to-GG splice-site variant in the intron 4 results in removal of the exon 4. The distributions of these non-shattering haplotypes among sorghum landraces suggest three independent origins. The function of the rice ortholog (OsSh1) was subsequently validated with a shattering-resistant mutant, and two maize orthologs (ZmSh1-1 and ZmSh1-5.1+ZmSh1-5.2) were verified with a large mapping population. Our results indicate that Sh1 genes for seed shattering were under parallel selection during sorghum, rice and maize domestication.

Plant domestication through an ecological lens

Abstract: Our understanding of domestication comes largely from archeology and genetics. Here, we advocate using current ecological theory and methodologies to provide novel insights into the causes and limitations of evolution under cultivation, as well as into the wider ecological impacts of domestication. We discuss the importance of natural selection under cultivation, that is, the forces promoting differences in Darwinian fitness between plants in crop populations and of constraints, that is, limitations of diverse nature that, given values for trait X, shorten the range of variation of trait Y, during the domestication process. Throughout this opinion paper, we highlight how ecology can yield insight into the effects of domestication on plant traits, on crop feedback over ecological processes, and on how species interactions develop in croplands.