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bioRxiv2017

A central role for boom-bust cycles in maintaining genetic variation in fluctuating environments

The long-running debate about the role of selection in maintaining genetic variation has been given new impetus by the discovery of hundreds of seasonally oscillating polymorphisms in wild Drosophila, possibly stabilized by an alternating summer-winter selection regime. Historically there has been skepticism about the potential of temporal variation to balance polymorphism, because selection must be strong to have a meaningful stabilizing effect -- unless dominance also varies over time ("reversal of dominance"). Here we develop a simplified model of seasonally variable selection that simultaneously incorporates four different stabilizing mechanisms, including two genetic mechanisms ("cumulative overdominance" and reversal of dominance), as well as ecological "storage" ("protection from selection" and boom-bust demography). We use our model to compare the stabilizing effects of these mechanisms. Although reversal of dominance has by far the greatest stabilizing effect, we argue that the three other mechanisms could also stabilize polymorphism under plausible conditions, particularly when all three are present. With many loci subject to diminishing returns epistasis, reversal of dominance stabilizes many alleles of small effect. This makes the combination of the other three mechanisms, which are incapable of stabilizing small effect alleles, a better candidate for stabilizing the detectable frequency oscillations of large effect alleles.

evolutionary biology↗

bioRxiv2017

Collaborative environmental DNA sampling from petal surfaces of flowering cherry Cerasus ×- yedoensis 'Somei-yoshino' across the Japanese archipelago

Recent studies have shown that environmental DNA is found almost everywhere. Flower petal surfaces are an attractive tissue to use for investigation of the dispersal of environmental DNA in nature as they are isolated from the external environment until the bud opens and only then can the petal surface accumulate environmental DNA. Here, we performed a crowdsourced experiment, the \"Ohanami Project\", to obtain environmental DNA samples from petal surfaces of Cerasus x yedoensis Somei-yoshino across the Japanese archipelago during spring 2015. C. x yedoensis is the most popular garden cherry species in Japan and clones of this cultivar bloom simultaneously every spring. Data collection spanned almost every prefecture and totaled 577 DNA samples from 149 collaborators. Preliminary amplicon-sequencing analysis showed the rapid attachment of environmental DNA onto the petal surfaces. Notably, we found DNA of other common plant species in samples obtained across a wide distribution; this DNA likely originated from pollen of the Japanese cedar. Our analysis supports our belief that petal surfaces after blossoming are a promising target to reveal the dynamics of environmental DNA in nature. The success of our experiment also shows that crowdsourced environmental DNA analyses have considerable value in ecological studies.

microbiology↗

bioRxiv2017

Fantastic beasts and how to sequence them: genomic approaches for obscure model organisms.

Application of genomic approaches to \"obscure model organisms\" (OMOs), meaning species with little or no genomic resources, enables increasingly sophisticated studies of genomic basis of evolution, acclimatization and adaptation in real ecological contexts. Here, I highlight sequencing solutions and data handling techniques most suited for genomic analysis of OMOs.\n\nGlossary- Allele Frequency Spectrum, AFS (same as Site Frequency Spectrum, SFS): histogram of the number of segregating variants depending on their frequency in one or more populations.\n- Restriction site-Associated DNA (RAD) sequencing: family of diverse genotyping methods that sequence short fragments of the genome adjacent to recognition site(s) for specific restriction endonuclease(s).\n- Linkage Disequilibrium (LD): in this review, correlation of genotypes at a pair of markers across individuals.\n- LD block: typical distance between markers in the genome across which their genotypes remain correlated.\n- Genome scan: profiling of genotypes along the genome looking for unusual patterns. Often used to look for signatures of natural selection or introgression.\n- \"Denser-than-LD\" genotyping: genotyping of several polymorphic markers per LD block.\n- Highly contiguous reference: genome or transcriptome reference sequence containing the least amount of fragmentation.\n- Phased data: data showing which SNP alleles belong to the same homologous chromosome copy.\n- Cross-tissue gene expression analysis: looking for individual-specific shifts in gene expression detectable across multiple tissues. Such shifts are predominantly genetic in nature.

genomics↗

bioRxiv2017

Cooperating yet distinct brain networks engaged during naturalistic paradigms: A meta-analysis of functional MRI results

Cognitive processes do not occur by pure insertion and instead depend on the full complement of co-occurring mental processes, including perceptual and motor functions. As such, there is limited ecological validity to human neuroimaging experiments that use highly controlled tasks to isolate mental processes of interest. However, a growing literature shows how dynamic, interactive tasks have allowed researchers to study cognition as it more naturally occurs. Collective analysis across such neuroimaging experiments may answer broader questions regarding how naturalistic cognition is biologically distributed throughout the brain. We applied an unbiased, data-driven, meta-analytic approach that uses k-means clustering to identify core brain networks engaged across the naturalistic functional neuroimaging literature. Functional decoding allowed us to, then, delineate how information is distributed between these networks throughout the execution of dynamical cognition in realistic settings. This analysis revealed seven recurrent patterns of brain activation, representing sensory, domain-specific, and attentional neural networks that support the cognitive demands of naturalistic paradigms. Though gaps in the literature remain, these results suggest that naturalistic fMRI paradigms recruit a common set of networks that that allow both separate processing of different streams of information and integration of relevant information to enable flexible cognition and complex behavior.

neuroscience↗

bioRxiv2017

Developmental transcriptomics of the brittle star Amphiura filiformis reveals gene regulatory network rewiring in echinoderm larval skeleton evolution

Amongst the echinoderms the class Ophiuroidea is of particular interest for its phylogenetic position, ecological importance, developmental and regenerative biology. However, compared to other echinoderms, notably echinoids (sea urchins), relatively little is known about developmental changes in gene expression in ophiuroids. To address this issue we have generated and assembled a large RNAseq data set of four key stages of development in the brittle star Amphiura filiformis and a de novo reference transcriptome of comparable quality to that of a model echinoderm - the sea urchin Strongyloncentrotus purpuratus. Furthermore, we provide access to the new data via a web interface: http://www.echinonet.eu/shiny/Amphiura_filiformis/. With a focus on skeleton development, we have identified highly conserved genes associated with the development of a biomineralized skeleton. We also identify important class-specific characters, including the independent duplication of the msp130 class of genes in different echinoderm classes and the unique occurrence of spicule matrix (sm) genes in echinoids. Using a new quantification pipeline for our de novo transcriptome, validated with other methodologies, we find major differences between brittle stars and sea urchins in the temporal expression of many transcription factor genes. This divergence in developmental regulatory states is more evident in early stages of development when cell specification begins, than when cells initiate differentiation. Our findings indicate that there has been a high degree of gene regulatory network rewiring in the evolution of echinoderm larval development.\n\nData DepositionsAll sequence reads are available at Genbank SRR4436669 - SRR4436674. Any sequence alignments used are available by the corresponding author upon request.

evolutionary biology↗

bioRxiv2017

The human auditory brainstem response to running speech reveals a subcortical mechanism for selective attention

Humans excel at selectively listening to a target speaker in background noise such as competing voices. While the encoding of speech in the auditory cortex is modulated by selective attention, it remains debated whether such modulation occurs already in subcortical auditory structures. Investigating the contribution of the human brainstem to attention has, in particular, been hindered by the tiny amplitude of the brainstem response. Its measurement normally requires a large number of repetitions of the same short sound stimuli, which may lead to a loss of attention and to neural adaptation. Here we develop a mathematical method to measure the auditory brainstem response to running speech, an acoustic stimulus that does not repeat and that has a high ecological validity. We employ this method to assess the brainstems activity when a subject listens to one of two competing speakers, and show that the brainstem response is consistently modulated by attention.

neuroscience↗

bioRxiv2017

An updated infrageneric classification of the oaks: review of previous taxonomic schemes and synthesis of evolutionary patterns

In this paper, we review major classification schemes proposed for oaks by John Claudius Loudon, Anders Sandoe Orsted, William Trelease, Otto Karl Anton Schwarz, Aimee Antoinette Camus, Yuri Leonardovich Menitsky, and Kevin C. Nixon. Classifications of oaks (Fig. 1) have thus far been based entirely on morphological characters. They differed profoundly from each other because each taxonomist gave a different weight to distinguishing characters; often characters that are homoplastic in oaks. With the advent of molecular phylogenetics our view has considerably changed. One of the most profound changes has been the realisation that the traditional split between the East Asian subtropical to tropical subgenus Cyclobalanopsis and the subgenus Quercus that includes all other oaks is artificial. The traditional concept has been replaced by that of two major clades, each comprising three infrageneric groups: a Palearctic-Indomalayan clade including Group Ilex (Ilex oaks), Group Cerris (Cerris oaks) and Group Cyclobalanopsis (cycle-cup oaks), and a predominantly Nearctic clade including Group Protobalanus (intermediate or golden cup oaks), Group Lobatae (red oaks) and Group Quercus (white oaks, with most species in America and some 30 species in Eurasia). The main morphological feature characterising these phylogenetic lineages is pollen morphology, a character overlooked in traditional classifications. This realisation, along with the now available (molecular-)phylogenetic framework, opens new avenues for biogeographic, ecological and evolutionary studies and a re-appraisal of the fossil record. We provide an overview about recent advances in these fields and outline how the results of these studies contribute to the establishment of a unifying systematic scheme of oaks. Ultimately, we propose an updated classification of Quercus recognising two subgenera with eight sections. This classification considers morphological traits, molecular-phylogenetic relationships, and the evolutionary history of one of the most important temperate woody plant genera.\n\nO_FIG O_LINKSMALLFIG WIDTH=156 HEIGHT=200 SRC=\"FIGDIR/small/168146_fig1.gif\" ALT=\"Figure 1\">\nView larger version (49K):\norg.highwire.dtl.DTLVardef@d5a05org.highwire.dtl.DTLVardef@91295forg.highwire.dtl.DTLVardef@3338fcorg.highwire.dtl.DTLVardef@e24a58_HPS_FORMAT_FIGEXP M_FIG O_FLOATNOFigure 1:C_FLOATNO Classification schemes for Quercus from Loudon to Nixon (see Appendix). Colour coding denotes the actual systematic affiliation of species included in each taxon: of the Old World or mid-latitude clade section Cyclobalanopsis (cycle-cup oaks, yellow), section Cerris (Cerris oaks; orange), and section Ilex (Ilex oaks; green); and of the New World or high-latitude clade section Quercus (white oaks s.str.; blue), sections Virentes (cyan) and Ponticae (dark blue), section Protobalanus (intermediate oaks; purple), and section Lobatae (red oaks; red). Colour gradients are proportional, i.e. reflect the proportion of species with different systematic affiliation included in each taxon. Note: Menitsky (1984) and Trelease (1924) only treated the Eurasian and American oaks, respectively, and provided classifications in (nearly) full agreement with current phylogenies.\n\nC_FIG

plant biology↗

bioRxiv2017

A GAL80 collection to nullify transgenes in Drosophila olfactory sensory neurons

Fruit flies recognize hundreds of ecologically relevant odors and respond appropriately to them. The complexity, redundancy and interconnectedness of the olfactory machinery complicate efforts to pinpoint the functional contributions of any component neuron or receptor to behavior. Some contributions can only be elucidated in flies that carry multiple mutations and transgenes, but the production of such flies is currently labor-intensive and time-consuming. Here, we describe a set of transgenic flies that express the Saccharomyces cerevisiae GAL80 in specific olfactory sensory neurons (OrX-GAL80s). The GAL80s effectively and specifically subtract the activities of GAL4-driven transgenes that impart anatomical and physiological phenotypes. OrX-GAL80s can allow researchers to efficiently activate only one or a few types of functional neurons in an otherwise nonfunctional olfactory background. Such experiments will improve our understanding of the mechanistic connections between odorant inputs and behavioral outputs at the resolution of only a few functional neurons.

neuroscience↗

bioRxiv2017

Effects of population- and seed bank noise on neutral evolution and efficacy of natural selection

Population genetics models typically consider a fixed population size and a unique selection coefficient. However, population dynamics inherently generate noise in numbers of individuals and selection acts on various components of the individuals fitness. In plant species with seed banks, the size of both the above- and below-ground compartments present noise depending on seed production and the state of the seed bank. We investigate if this noise has consequences on 1) the rate of genetic drift, and 2) the efficacy of selection. We consider four variants of two-allele Moran-type models defined by combinations of presence and absence of noise in above-ground and seed bank compartments. Time scale analysis and dimension reduction methods allow us to reduce the corresponding Fokker-Planck equation to a one-dimensional diffusive Moran model. We first show that if the above-ground noise classically affects the rate of genetic drift, below-ground noise reduces the diversity storage effect of the seed bank. Second, we consider that selection can act on four different components of the plant fitness: plant or seed death rate, seed production or seed germination. Our striking result is that the efficacy of selection for seed death rate or germination rate is reduced by seed bank noise, whereas selection occurring on plant death rate or seed production is not affected. We derive the expected site-frequency spectrum reflecting this heterogeneity in selection efficacy between genes underpinning different plant fitness components. Our results highlight the importance to consider the effect of ecological noise to predict the impact of seed banks on neutral and selective evolution.

evolutionary biology↗

bioRxiv2017

Facilitation of motor excitability during listening to spoken sentences is not modulated by noise or semantic coherence

Comprehending speech can be particularly challenging in a noisy environment and in the absence of semantic context. It has been proposed that the articulatory motor system would be recruited especially in difficult listening conditions. However, it remains unknown how signal-to-noise ratio (SNR) and semantic context affect the recruitment of the articulatory motor system when listening to continuous speech. The aim of the present study was to address the hypothesis that involvement of the articulatory motor cortex increases when the intelligibility and clarity of the spoken sentences decreases, because of noise and the lack of semantic context. We applied Transcranial Magnetic Stimulation (TMS) to the lip and hand representations in the primary motor cortex and measured motor evoked potentials from the lip and hand muscles, respectively, to evaluate motor excitability when young adults listened to sentences. In Experiment 1, we found that the excitability of the lip motor cortex was facilitated during listening to both semantically anomalous and coherent sentences in noise, but neither SNR nor semantic context modulated the facilitation. In Experiment 2, we replicated these findings and found no difference in the excitability of the lip motor cortex between sentences in noise and clear sentences without noise. Thus, our results show that the articulatory motor cortex is involved in speech processing even in optimal and ecologically valid listening conditions and that its involvement is not modulated by the intelligibility and clarity of speech.

neuroscience↗

bioRxiv2017

Double-digest RAD sequencing outperforms microsatellite loci at assigning paternity and estimating relatedness: a proof of concept in a highly promiscuous bird

Information on genetic relationships among individuals is essential to many studies of the behavior and ecology of wild organisms. Parentage and relatedness assays based on large numbers of SNP loci hold substantial advantages over the microsatellite markers traditionally used for these purposes. We present a double-digest restriction site-associated DNA sequencing (ddRAD-seq) analysis pipeline that, as such, simultaneously achieves the SNP discovery and genotyping steps and which is optimized to return a statistically powerful set of SNP markers (typically 150-600 after stringent filtering) from large numbers of individuals (up to 240 per run). We explore the tradeoffs inherent in this approach through a set of experiments in a species with a complex social system, the variegated fairy-wren (Malurus lamberti), and further validate it in a phylogenetically broad set of other bird species. Through direct comparisons with a parallel dataset from a robust panel of highly variable microsatellite markers, we show that this ddRAD-seq approach results in substantially improved power to discriminate among potential relatives and considerably more precise estimates of relatedness coefficients. The pipeline is designed to be universally applicable to all bird species (and with minor modifications to many other taxa), to be cost- and time-efficient, and to be replicable across independent runs such that genotype data from different study periods can be combined and analyzed as field samples are accumulated.

animal behavior and cognition↗

bioRxiv2017

Bringing plants and soils to life through a simple role-playing activity

Interactions are at the core of many ecological and evolutionary forces in nature. Plant-soil interactions provide a rich example of the interconnectedness of living systems, but are hidden from everyday view and overshadowed in the classroom by more popular teaching examples involving animals, reptiles, or invertebrates. To highlight the importance and relevance of plant-soil relationships, we devised a simple role-playing activity suitable for college students. Specifically, the activity simulates how feedbacks between plants and soil environments influence plant species abundance and community richness. With this activity, students will gain a better understanding of these prolific, but overlooked, forms of biological interactions that impact the diversity and functioning of ecosystems.

scientific communication and education↗

bioRxiv2017

Response of oriental xerophytes to the occidental industrial revolution

Since the Industrial Revolution, human activities have contributed substantially to climate change, by adding CO2 to the atmosphere, especially since the mid-20th century (the \"Great Acceleration\"). Climatic change does not have the same impact on different regions of the Earth, neither in the recorded past, nor in models of the future. Therefore, to anticipate on these changes, we need to understand and be able to predict the possible responses of the different regional vegetations of the world to these changes, and most significantly to increased drought conditions. The aim of this study is to understand the response of the xerophytes of the immense oriental Irano-Turanian bioregion to post-industrial global warming and to compare it with the response of the xerophytes of the neighbouring occidental Mediterranean bioregion. We measured stomata index and stomata density from 83 herbarium sheets (coll. 1821-2014) from species of the non-succulent xerophyte, Haplophyllum. We tested for differences before and after the \"Great Acceleration\" in both bioregions. SI decreased in the occidental species (significant only for abaxial leaf side), whereas SI significantly increased in the oriental species (both sides). We suggest that changes in both occidental and oriental species are linked to atmospheric CO2 due to the different constraints that act on their growth. In light-limited occidental species, atmospheric CO2 caused the stomata index decrease, whereas in the predominantly water-limited oriental species, increased drought stress and temperature (climatic change) caused stomata index increase. In conclusion, we propose that whereas atmospheric CO2 directly caused a decrease in stomata index in occidental xerophytes, it indirectly caused an increase in stomata index in oriental xerophytes via climate change (increase in aridity, drought stress, and temperature). This study highlights the considerable potential of research based on historical herbarium collections to answer ecological questions, especially regarding climatic change.

plant biology↗

bioRxiv2017

The genetic architecture of the network underlying flowering time variation in Arabidopsis thaliana

Flowering time is a key adaptive trait in plants and is tightly controlled by a complex regulatory network that responds to seasonal signals. In a rapidly changing climate, understanding the genetic basis of flowering time variation is important for both agriculture and ecology. Genetic mapping has revealed many genetic variants affecting flowering time, but the effects on the gene regulatory networks in population-scale are still largely unknown. We dissected flowering time networks using multi-layered Swedish population data from Arabidopsis thaliana, consisting of flowering time and transcriptome collected under constant 10{degrees}C growth temperature in addition to full genome sequence data. Our analysis identified multiple alleles of the key flowering time gene FLOWERING LOCUS C (FLC) as the primary determinant of the network underlying flowering time variation under our condition. Genetic variation of FLC affects multiple-pathways through known flowering-time genes including FLOWERING LOCUS T (FT), and SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1). We demonstrated that an extremely simple single-locus model of FLC involving allelic variation and expression explains almost a half of flowering time variation, with 60% of the effect being mediated through FLC expression. Furthermore, the accuracy of the model fitted at 10{degrees}C is almost unchanged at 16{degrees}C.

genetics↗

bioRxiv2017

A family-based method for leveraging random genetic variation to identify variance-controlling loci

The propensity of a trait to vary within a population may have evolutionary, ecological, or clinical significance. In the present study we deploy sibling models to offer a novel and unbiased way to ascertain loci associated with the extent to which phenotypes vary (variance-controlling quantitative trait loci, or vQTLs). Previous methods for vQTL-mapping either exclude genetically related individuals or treat genetic relatedness among individuals as a complicating factor addressed by adjusting estimates for non-independence in phenotypes. The present method uses genetic relatedness as a tool to obtain unbiased estimates of variance effects rather than as a nuisance. The family-based approach, which utilizes random variation between siblings in minor allele counts at a locus, also allows controls for parental genotype, mean effects, and non-linear (dominance) effects that may spuriously appear to generate variation.\n\nSimulations show that the approach performs equally well as two existing methods (squared Z-score and DGLM) in controlling type I error rates when there is no unobserved confounding, and performs significantly better than these methods in the presence of confounding. Using height and BMI as empirical applications, we investigate SNPs that alter within-family variation in height and BMI, as well as pathways that appear to be enriched. One significant SNP for BMI variability, in the MAST4 gene, replicated. Pathway analysis revealed one gene set, encoding members of several signaling pathways related to gap junction function, which appears significantly enriched for associations with within-family height variation in both datasets (while not enriched in analysis of mean levels). We recommend approximating laboratory random assignment of genotype using family data and more careful attention to the possible conflation of mean and variance effects.

genetics↗

bioRxiv2017

Continuous gene flow contributes to low global species abundance and distribution of a marine model diatom

Diatoms emerged in the Mesozoic period and presently constitute one of the main primary producers in the worlds ocean and are of a major economic importance. In the current study, using whole genome sequencing of ten accessions of the model diatom Phaeodactylum tricornutum, sampled at broad geospatial and temporal scales, we draw a comprehensive landscape of the genomic diversity within the species. We describe strong genetic subdivisions of the accessions into four genetic clades (A-D) with constituent populations of each clade possessing a conserved genetic and functional makeup, likely a consequence of the limited dispersal of P. tricornutum in the open ocean. We further suggest dominance of asexual reproduction across all the populations, as implied by high linkage disequilibrium. Finally, we show limited yet compelling signatures of genetic and functional convergence inducing changes in the selection pressure on many genes and metabolic pathways. We propose these findings to have significant implications for understanding the genetic structure of diatom populations in nature and provide a framework to assess the genomic underpinnings of their ecological success and impact on aquatic ecosystems where they play a major role. Our work provides valuable resources for functional genomics and for exploiting the biotechnological potential of this model diatom species.

genomics↗

bioRxiv2017

Horizontal gene cluster transfer increased hallucinogenic mushroom diversity

Secondary metabolites are heterogeneous natural products that often mediate interactions between species. The tryptophan-derived secondary metabolite, psilocin, is a serotonin receptor agonist that induces altered states of consciousness. A phylogenetically disjunct group of mushroom-forming fungi in the Agaricales produce the psilocin prodrug, psilocybin. Spotty phylogenetic distributions of fungal compounds are sometimes explained by horizontal transfer of metabolic gene clusters among unrelated fungi with overlapping niches. We report the discovery of a psilocybin gene cluster in three hallucinogenic mushroom genomes, and evidence for its horizontal transfer between fungal lineages. Patterns of gene distribution and transmission suggest that psilocybin provides a fitness advantage in the dung and late wood-decay niches, which may be reservoirs of fungal indole-based metabolites that alter behavior of mycophagous and wood-eating invertebrates. These hallucinogenic mushroom genomes will serve as models in neurochemical ecology, advancing the prospecting and synthetic biology of novel neuropharmaceuticals.

evolutionary biology↗

bioRxiv2017

The Complex Ecosystem in Non Small Cell Lung Cancer Invasion

Many tumors are characterized by genetic instability, producing an assortment of genetic variants of tumor cells called subclones. These tumors and their surrounding environments form complex multi-cellular ecosystems, where subclones compete for resources and cooperate to perform multiple tasks, including cancer invasion. Our recent empirical studies revealed existence of such distinct phenotypes of cancer cells, leaders and followers, in lung cancer. These two cellular subclones exchange a complex array of extracellular signals demonstrating a symbiotic relationship at the cellular level. Here, we develop a computational model of the microenvironment of the lung cancer ecosystem to explore how the interactions between subclones can advance or inhibit invasion. We found that, due to the complexity of the ecosystem, invasion may have very different dynamics characterized by the different levels of aggressiveness. By altering the signaling environment, we could alter the ecological relationship between the cell types and the overall ecosystem development. Competition between leader and follower cell populations (defined by the limited amount of resources), positive feedback within the leader cell population (controlled by the focal adhesion kinase and fibronectin signaling), and impact of the follower cells to the leaders (represented by yet undetermined proliferation signal) all had major effects on the outcome of the collective dynamics. Specifically, our analysis revealed a class of tumors (defined by the strengths of fibronectin signaling and competition) that are particularly sensitive to manipulations of the signaling environment. This class can undergo irreversible changes to the tumor ecosystem that outlast these manipulations of feedbacks and have a profound impact on invasive potential. Our study predicts a complex division of labor between cancer cell subclones and suggests new treatment strategies targeting signaling within the tumor ecosystem.\n\nAuthor SummaryCancer is an elusive disease due to the wide variety of cancer types and adaptability to treatment. How is this adaptability accomplished? Loss of genetic stability, a hallmark of cancer, leads to the emergence of many different types of cancer cells within a tumor. This creates a complex ecosystem where cancer cell types can cooperate, compete, and exploit each other. We have previously used an image-guided technology to isolatedistinct cancer subclones and identify how they interact. Here, we have employed mathematical modeling to understand how the dynamic feedbacks between different cancer cell types can impact the success of invasion in lung cancer. We found that successful invasion required for feedbacks to support the less viable but more invasive cell types. These predictions may have implications for novel clinical treatment options and emphasize the need to visualize and probe cancer as a tumor ecosystem.

cancer biology↗