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Velicer, G. J.

Publications and source records attributed to Velicer, G. J..

14 recordsLinked to original sources
bioRxiv2026

CRISPR-Cas interference decays rapidly with distance from the leader sequence in a long array

Spacer efficacy generally declines with distance from the leader sequence, but the scarcity of fine-scale studies hampers comparisons across taxa. Here, we investigated positional effects across an exceptionally long 121-spacer CRISPR array associated with the type I-C cas operon of a Myxococcus xanthus natural isolate. In plasmid-interference assays, we found that interference rapidly declined with distance from the leader sequence, with only the proximal [~]4% of spacers conferring measurable interference. This contrasts strikingly with a study in Vibrio cholerae, in which it was shown that [~]95% of spacers in a shorter (39-spacer) array were effective. Our results suggest that there is great variation in the effective proportion of spacers across species, highlighting the need for fine-scale studies of CRISPR-array activity across diverse bacterial lineages.

microbiology↗
bioRxiv2026

Predator adaptation to single prey species yields positive specialization and multiple forms of diversification

Ecological specialization emerges when adaptation to a focal context increases fitness in that context relative to others. Experimental evolution has been widely used to study microbial specialization in abiotic environments but not predatory specialization. Here we demonstrate evolutionary specialization by a bacterial predator and characterize associated diversification of its predation profile. Populations of the bacterium Myxococcus xanthus evolving on single, non-evolving prey species diverged in predatory performance, showing increased performance on their home prey relative to their common ancestor, and relative to foreign prey not encountered during adaptation. Adaptation to the single-prey environments resulted in striking radiation of performance profiles across a diverse panel of foreign prey that was shaped interactively by selection, chance, and indirect effects, with home-prey identity modulating the degree of stochastic indirect diversification. Despite a great diversity of indirect evolutionary effects, correlated evolution was net-positive, yielding positive predatory specialization as the general outcome. Genomic evolution mirrored phenotypic evolution in that degrees of genomic parallelism differed as a function of home-prey identity. These findings show that adaptation to even simple biotic conditions can generate great ecological and behavioral diversity, linking direct selection, deterministic indirect effects of adaptation, stochasticity, and the origins of predator specialization and diversification.

evolutionary biology↗
bioRxiv2025

Predation, evo-devo, and historical contingency: A nematode predator drives evolution of aggregative multicellularity

Regulated aggregation and differentiation of cells is an intriguing form of multicellularity that produces diverse morphological forms, resulting from an interplay of complex evolutionary, regulatory, behavioral, and social mechanisms. We investigate the potential for predation to shape aggregative multicellularity evolution across different combinations of three species comprising a multi-trophic food web. The bacterium Myxococcus xanthus, which can develop into spore-filled fruiting bodies, is a mesopredator; the bacterivorous nematode Pristionchus pacificus is an apex predator, and the bacterium Escherichia coli is a shared basal prey for both predators. The number and morphology of M. xanthus fruiting bodies responds evolutionarily to nematodes, regardless of whether E. coli is present. E. coli alone with M. xanthus reduces both fruiting body formation and spore production, but the presence of nematodes eliminates those effects. M. xanthus lineages with an ancestral antibiotic resistance mutation evolved less overall, revealing strong historical contingency and suggesting potential tradeoffs between antibiotic resistance and responsiveness to biotic selection. Our results suggest that predation both of and by mesopredators has played important roles in the evolution of aggregative multicellularity and reveal complex inter-trophic evolutionary interactions in a relatively simple three-species food web.

evolutionary biology↗
bioRxiv2024

Deletion of an sRNA primes development in a multicellular bacterium

ABSTRACTSmall non-coding RNAs (sRNAs) are essential in regulating gene expression during many biological processes. The myxobacteria gene pxr encodes an sRNA known to block fruiting-body development, an aggregative multicellular process triggered by starvation. Deletion of pxr allows Myxococcus xanthus cells to develop in the presence of nutrients. However, potential Pxr binding targets and most genes regulated by Pxr remain unknown. Here, we found that the absence of pxr expression dramatically alters the temporal dynamics of development, thus suggesting an important new role of this sRNA in myxobacterial ecology. We transcriptionally profiled vegetative cells of M. xanthus strains possessing vs lacking pxr and found that over half of the genes impacted by pxr deletion during growth are linked to development, including known and potentially novel critical regulators. Many other genes are associated with general metabolic processes, which Pxr regulates positively. Our study discovers new phenotypic effects of Pxr regulation of likely ecological importance, identifies the suite of genes this sRNA controls during vegetative growth, reveals a previously unknown developmental regulator and provides new insights into the early molecular regulation of myxobacterial development.

molecular biology↗
bioRxiv2023

Ecological histories determine the success of social exploitation

Ecological context often modifies biotic interactions, yet effects of ecological history are poorly understood. In experiments with the bacterium Myxococcus xanthus, resource-level histories of genotypes interacting during cooperative multicellular development were found to strongly regulate social fitness. Yet how developmental spore production responded to variation in resource-level histories between interactants differed greatly between cooperators and cheaters; relative-fitness advantages gained by cheating after high-resource growth were generally reduced or absent if one or both parties experienced low-resource growth. Low-resource growth also eliminated facultative exploitation in some pairwise mixes of cooperation-proficient natural isolates that occurs when both strains have grown under resource abundance. Our results contrast with previous studies in which cooperator fitness correlated positively with resource level and suggest that resource-level variation may be important in regulating whether exploitation of cooperators occurs in a natural context.

ecology↗
bioRxiv2022

Antagonistic, synergistic, and social pleiotropy in microbial cheaters

Cooperation is widespread among microbes. One mechanism proposed to constrain cheating is antagonistic pleiotropy, wherein mutations that cause defection from cooperation, while potentially under positive selection for this effect, reduce fitness at other traits. The bacterium Myxococcus xanthus engages in pleiotropically connected complex multicellular behaviors, including motility, predation, and starvation-induced fruiting-body development. Sporulation during development is susceptible to cheating. Here we investigate pleiotropic impacts on how cheater spores respond to stressful environmental changes after development, relative to cooperator spores. A cheater with a mutation in the developmental-signaling gene asgB shows antagonistic pleiotropy under both heat and basic-pH stress, while a cheater mutated at csgA shows synergistic pleiotropy under basic-pH stress. Further, in a social form of pleiotropy, cooperator spores formed in mixture with the asgB cheater are less resilient under basic pH than those from pure groups; interaction of cooperators with cheaters reduces the cooperators physical robustness. Our results indicate that, depending on the mutation, pleiotropy can promote as well as limit cheating alleles. They additionally demonstrate that alleles can pleiotropically alter traits in organisms not carrying those alleles. Synergistic and social pleiotropy may contribute to shaping the evolutionary dynamics of cooperation and cheating in many social systems.

evolutionary biology↗
bioRxiv2021

Widespread density dependence of bacterial growth under acid stress

Benefits of cooperation intrinsically depend on density because biological interaction requires organismal proximity. Microbial cooperative traits are common, yet systematic tests for a shared cooperative phenotype across diverse species are rare, as are direct tests for the Allee effect - positive density dependence of fitness. Here we test for positive density dependence of growth under acid stress in five phylogenetically widespread bacterial species - three Gram-negative and two Gram-positive - and find the Allee effect in all five. However, social protection from acid stress appears to have evolved by different mechanisms across species. In Myxococcus xanthus, the acid-stress Allee effect is found to be mediated by pH-regulated secretion of a diffusible molecule present in supernatant from high-density cultures. In contrast, growth from low density under acid stress by the other species was not enhanced by high-density supernatant. Additionally, density dependence of Myxococcus fruiting-body formation during starvation is found to increase with acid stress, suggesting that abiotic stresses other than starvation shape the evolution of aggregative development. Our findings suggest that high cell density may protect against acid stress in most bacterial species and in Myxococcus may promote predation on microbes that acidify their local environment by secretion of metabolic byproducts.

microbiology↗
bioRxiv2021

Hidden paths to endless forms most wonderful: Ecology latently shapes evolution of multicellular development in predatory bacteria

Ecological causes of developmental-system evolution, for example from predation, remain under intense investigation. An important open question is the role of latent phenotypes in eco-evo-devo. The predatory bacterium Myxococcus xanthus undergoes aggregative multicellular development upon starvation. Here we use M. xanthus to test whether evolution in several distinct growth environments that do not induce development latently alters developmental phenotypes, including morphology and plasticity, in environments that do induce development. In the MyxoEE-3 evolution experiment, growing M. xanthus populations swarmed across agar surfaces while adapting to distinct conditions varying at factors such as surface stiffness or prey identity. All examined developmental phenotypes underwent extensive and ecologically specific latent evolution, with surface stiffness, prey presence and prey identity all strongly impacting the latent evolution of development. Evolution on hard agar allowed retention of developmental proficiency and extensive stochastic phenotypic radiation, including of reaction norms, with instances of both increased plasticity and canalization. In contrast, evolution on soft agar latently led to systematic loss of development, revealing an ecologically-contingent fitness trade-off between the growth and developmental phases of a multicellular life cycle that is likely determined by details of motility behavior. Similar contingency was observed after evolution during predatory growth in distinct prey environments, with Bacillus subtilis causing greater loss of development and lower stochastic diversification than Escherichia coli. Our results have implications for understanding evolutionary interactions among predation, development and motility in myxobacterial life cycles, and, more broadly, the importance of latent phenotypes for the diversification of developmental systems.

evolutionary biology↗
bioRxiv2021

Social selection within aggregative multicellular development drives morphological evolution

The evolution of developmental systems might be shaped by both historical differences in developmental features and social selection, among other factors. In aggregative multicellularity, development is itself a social process in which unicellular organisms cooperate in carrying out complex developmental programs. In some aggregative systems, development culminates in the construction of spore-packed fruiting bodies. Fruiting body development in myxobacteria often unfolds within genetically and behaviorally diverse conspecific cellular environments that can include social defection and warfare. Here we use the bacterium Myxococcus xanthus to test whether the character of the cellular environment during aggregative development shapes morphological evolution. We manipulated the cellular composition of Myxococcus development in an experiment in which evolving populations initiated from a single ancestor repeatedly co-developed with one of several non-evolving partners - a benign cooperator, one of three cheaters or one of three antagonists. Fruiting body morphology was found to diversify as a function of developmental partners, revealing adaptation specific to distinct cellular environments. Collectively, antagonistic partners selected for higher levels of robust fruiting body formation than did cheaters or the benign cooperator. Moreover, even small degrees of genetic divergence between the distinct cheater partners were sufficient to drive treatment-level morphological divergence. Co-developmental partners not only shaped mean trait evolution but also determined the magnitude and dynamics of stochastic morphological diversification and subsequent convergence. In sum, we find that even few genetic differences affecting developmental and social features can greatly impact the morphological evolution of multicellular bodies and experimentally demonstrate that microbial warfare can promote cooperation.

evolutionary biology↗
bioRxiv2021

Behavioral interactions between bacterivorous nematodes and predatory bacteria in a synthetic community

Theory and empirical studies in metazoans predict that apex predators should shape the behavior and ecology of mesopredators and prey at lower trophic levels. Despite the ecological importance of microbial communities, few studies of predatory microbes examine such behavioral responses and the multiplicity of trophic interactions. Here, we sought to assemble a three-level microbial food chain and to test for behavioral interactions between the predatory nematode Caenorhabditis elegans and the predatory social bacterium Myxococcus xanthus when cultured together with two basal prey bacteria that both predators can eat - Escherichia coli and Flavobacterium johnsoniae. We find that >90% of C. elegans worms failed to interact with M. xanthus even when it was the only potential prey species available, whereas most worms were attracted to pure patches of E. coli and F. johnsoniae. In addition, M. xanthus altered nematode predatory behavior on basal prey, repelling C. elegans from two-species patches that would be attractive without M. xanthus, an effect similar to that of C. elegans pathogens. The nematode also influenced the behavior of the bacterial predator: M. xanthus increased its predatory swarming rate in response to C. elegans in a manner dependent both on basal-prey identity and on worm density. Our results suggest that M. xanthus is an unattractive prey for some soil nematodes and is actively avoided when more lucrative prey are available. Most broadly, we find that nematode and bacterial predators mutually influence one anothers predatory behavior, with likely consequences for coevolution within complex microbial food webs.

ecology↗
bioRxiv2021

Allopatric divergence limits cheating range and alters genetic requirements for a cooperative trait

Social defectors may meet diverse cooperators. Genotype-by-genotype interactions may constrain the ranges of cooperators upon which particular defectors can cheat, limiting cheater spread. The bacterium Myxococcus xanthus undergoes cooperative multicellular development, but some developmental defectors cheat on cooperators during this process. In this study, interactions between a cheater disrupted at the signaling gene csgA and allopatrically diversified cooperators reveal a very small cheating range. Expectedly, the cheater failed to cheat on all natural-isolate cooperators owing to non-cheater-specific antagonisms. Surprisingly, lab-evolved cooperators that diverged from their cheating-susceptible ancestor by fewer than 20 mutations and without experiencing cheating had already exited the csgA mutants cheating range. Cooperators might also diversify in the potential for a mutation to reduce expression of cooperative trait or generate a cheating phenotype. A new csgA mutation constructed in several highly diverged cooperators generated diverse sporulation phenotypes, ranging from a complete defect to no defect, indicating that genetic backgrounds can limit the set of genomes in which a mutation creates a defector. Our results suggest that natural populations feature geographic mosaics of cooperators that have diversified in their susceptibility to particular cheaters and in the phenotypes generated by any given cooperation-gene mutation. Significance statementSelection on cooperators exploited by obligate cheaters can induce evolution of resistance to cheating. Here we show that cooperators can also rapidly evolve immunity to cheating simply as a byproduct of evolutionary divergence in environments in which cooperation and cheating at the focal trait do not occur because the trait is not expressed. We also find that differences in the genomic context in which a cooperation-gene mutation arises can profoundly alter its phenotypic effect and determine whether the mutation generates a social defect at all - a pre-requisite for obligate cheating. These findings suggest that general divergence of social populations under a broad range of environmental conditions can restrict both the set of mutations that might generate social defectors in the first place and the host range of such defectors once they arise.

evolutionary biology↗
bioRxiv2020

Hidden paths to endless forms most wonderful: Parasite-blind diversification of host quality

Evolutionary diversification can occur in allopatry or sympatry, can be unselected or driven by selection, and can be phenotypically manifested immediately or remain phenotypically latent until later manifestation in a newly encountered environment. Diversification of host-parasite interactions is frequently studied in the context of intrinsically selective coevolution, but the potential for host-parasite interaction phenotypes to diversify latently during parasite-blind evolution is rarely considered. Here we use a social bacterium experimentally adapted to several environments in the absence of phage to analyse allopatric diversification of latent host quality - the degree to which a host population supports a viral epidemic. Phage-blind evolution reduced host quality overall, with some bacteria becoming completely resistant to growth suppression by phage. Selective-environment differences generated only mild divergence in host-quality. However, selective environments nonetheless played a major role in shaping evolution by determining the degree of stochastic diversification among replicate populations within treatments. Ancestral motility genotype was also found to strongly shape patterns of latent hostquality evolution and diversification. These outcomes show that adaptive landscapes can differ in how they constrain stochastic diversification of a latent phenotype and that major effects of selection on biological diversification can be missed by focusing on trait means. Collectively, our findings suggest that latent-phenotype evolution (LPE) should inform host-parasite evolution theory and that diversification should be conceived broadly to include latent phenotypes.

evolutionary biology↗
bioRxiv2019

Facultative social exploitation among Myxoccocus bacteria due to non-responsive local competition

Microbes cooperate in many ways, but it is unclear to what extent they also have adaptations to detect and exploit unrelated social partners. Experimental data often cannot discriminate between social exploitation caused by complex adaptive traits and exploitation caused by simple deleterious mutations. Here we demonstrate facultative social exploitation among Myxococcus bacteria due to simple non-responsive local competition. We show that the time dynamics of developmental exploitation by a tan phase variant are consistent with a model where cells do not respond to the presence of other genotypes but simply compete for some shared resource necessary to produce spores. The model also predicts the frequency-dependent responses of strains to genetic chimerism in multicellular fruiting bodies. Interactions between naturally occuring soil isolates, on the other hand, are consistent with strong interference competition where the winner almost completely prevents the loser from producing any spores. These results show how facultative social exploitation does not require elaborate mechanisms to detect and respond to foreign genotypes but can instead be caused by simple competition acting on a local scale. Myxococcus cells, like social insects, cooperate in some ways while simultaneously competing in others.

evolutionary biology↗