Horizontal, but not vertical, biotic interactions affect fine-scale plant distribution patterns in a low-energy system

Studies of species range determinants have traditionally focused on abiotic variables (typically climatic conditions), and therefore the recent explicit consideration of biotic interactions represents an important advance in the field. While these studies clearly support the role of biotic interactions in shaping species distributions, most examine only the influence of a single species and/or a single interaction, failing to account for species being subject to multiple concurrent interactions. By fitting species distribution models (SDMs), we examine the influence of multiple vertical (i.e., grazing, trampling, and manuring by mammalian herbivores) and horizontal (i.e., competition and facilitation; estimated from the cover of dominant plant species) interspecific interactions on the occurrence and cover of 41 alpine tundra plant species. Adding plant-plant interactions to baseline SDMs (using five field-quantified abiotic variables) significantly improved models' predictive power for independent data, while herbivore-related variables had only a weak influence. Overall, abiotic variables had the strongest individual contributions to the distribution of alpine tundra plants, with the importance of horizontal interaction variables exceeding that of vertical interaction variables. These results were consistent across three modeling techniques, for both species occurrence and cover, demonstrating the pattern to be robust. Thus, the explicit consideration of multiple biotic interactions reveals that plant-plant interactions exert control over the fine-scale distribution of vascular species that is comparable to abiotic drivers and considerably stronger than herbivores in this low-energy system.

Describing ancient horizontal gene transfers at the nucleotide and gene levels by comparative pathogenicity island genometrics.

MOTIVATION: Lateral gene transfer is a major mechanism contributing to bacterial genome dynamics and pathovar emergence via pathogenicity island (PAI) spreading. However, since few of these genomic exchanges are experimentally reproducible, it is difficult to establish evolutionary scenarios for the successive PAI transmissions between bacterial genera. Methods initially developed at the gene and/or nucleotide level for genomics, i.e. comparisons of concatenated sequences, ortholog frequency, gene order or dinucleotide usage, were combined and applied here to homologous PAIs: we call this approach comparative PAI genometrics. RESULTS: YAPI, a Yersinia PAI, and related islands were compared with measure evolutionary relationships between related modules. Through use of our genometric approach designed for tracking codon usage adaptation and gene phylogeny, an ancient inter-genus PAI transfer was oriented for the first time by characterizing the genomic environment in which the ancestral island emerged and its subsequent transfers to other bacterial genera.

Excision of the Staphylococcal Cassette Chromosome mec (SCCmec) and its crucial role in the horizontal transfer of methicillin resistance in staphylococci.

Staphylococcus aureus est un pathogène humain majeur ayant développé des résistances contre la quasi totalité des antibiotiques disponibles, incluant la très importante famille des β- lactamines. La résistance à cette classe d'antibiotiques est conférée par la « Staphylococcal Cassette Chromosome mec » (SCCmec), qui est un élément génétique mobile capable de s'insérer dans le chromosome bactérien et capable d'être transféré horizontalement chez d'autres staphylocoques. Le mécanisme moléculaire impliqué dans ce transfert horizontal demeure largement inconnu. L'une des premières étapes du transfert est l'excision du SCC mec du chromosome bactérien. Cette excision est promue par des enzymes codées par l'élément SCCmec lui- même et appelées de ce fait « Cassette Chromosome Recombinases » (Ccr). L'un des buts de ce travail de thèse a été de comprendre la régulation de l'expression des gènes codant pour les Ccr recombinases. En utilisant des outils moléculaires originaux, nous avons été en mesure de démontrer en premier lieu que les Ccr recombinases étaient exprimées de façon « bistable », c'est à dire qu'uniquement quelques pourcents de cellules dans une population exprimaient ces gènes à un temps donné. Dans un deuxième temps, nous avons également démontré que l'expression de ces gènes était régulée par des facteurs étrangers au SCC mec. L'expression bistable des recombinases est un concept important. Effectivement, cela permet à la majorité des cellules d'une population de conserver l'élément SCC mec, alors que seulement une petite fraction le perd afin de le rendre disponible pour un transfert. Ainsi, alors que l'élément SCC mec continue de se propager avec la multiplication des bactéries Staphylococcus aureus résistant à la méticilline (SARM), il peut être simultanément transmis à des souches susceptibles (Staphylococcus aureus susceptible à la méticilline, SASM), entraînant l'apparition de nouveaux SARM. De façon très intéressante, le fait que cette bistabilité est contrôlée par les bactéries, et non le SCCmec lui-même, montre que la décision de transférer ou non la cassette SCC mec appartient à la bactérie. En conséquence, il doit exister dans la nature des souches qui sont plus ou moins aptes à effectuer ce transfert. En nous appuyant sur ces observations, nous avons montré que l'excision du SCC mec était effectivement régulée de façon très étroite au cours de la division cellulaire, et ne se passait que pendant un temps limité au début de la croissance. Ce résultat est compatible avec une régulation génétique commandée par la densité cellulaire, qui pourrait être dépendante de la production de signaux extracellulaires, du type que l'on rencontre dans le quorum sensing. Les signaux hypothétiques entraînant l'excision du SCC mec restent inconnus à l'heure actuelle. La connaissance de ces signaux pourrait se révéler très importante afin de développer des stratégies pour interférer avec la dissémination de la résistance au β-lactamines. Deux sujets additionnels ont été logiquement investigués au vu de ces premiers résultats. Premièrement, si certaines souches de SARM sont plus ou moins aptes à déclencher l'excision du SCC mec, de même certaines souches de SASM devraient être plus ou moins aptes à acquérir cet élément. Deuxièmement, afin d'étudier ces mécanismes de transfert au niveau épidémiologique, il nous a été nécessaire de développer des outils nous permettant d'explorer le phénomène à une plus large échelle. Concernant le premier point, il a été postulé que certains SASM seraient réfractaires à l'intégration génomique d'un SCC mec en raison de polymorphismes particuliers à proximité du site d'insertion chromosomique (attB). En étudiant plus de 40 isolais de S. aureus, provenant de porteurs sains, nous avons confirmé ce polymorphisme dans l'environnement à'attB. De plus, nous avons pu montrer que ces régions polymorphiques ont évolué parallèlement à des groupes phylogénétiques bien connus. Ainsi, si des telles régions réfractaires à l'intégration de SCC mec existent, celles-ci devraient ségréger dans des complexes clonaux bien définis qui devraient être facilement identifiables au niveau épidémiologique. Concernant le second point, nous avons été capables de construire un système rapporteur de l'excision du SCCmec, en utilisant un plasmide à faible copie. Ce système consistait en un promoteur fort et un gène codant pour une protéine verte fluorescente (GFP) sous le contrôle d'un promoteur fort séparés à l'aide d'un élément SCC artificiel portant trois terminateurs de transcription. Ainsi, la fluorescence ne s'exprime que si l'élément SCC est excisé du plasmide. Ce système a été testé avec succès dans plusieurs types de staphylocoques, et est actuellement évalué dans d'autres souches et conditions stimulant ou inhibant l'excision. De manière générale, cette dissertation représente parcours scientifique à travers plusieurs aspects d'un problème de santé publique majeur en rapport avec la résistance bactérienne aux antibiotiques. Ce travail s'attaque à des problèmes fondamentaux concernant le transfert horizontal de l'élément SCC mec. De plus, il s'intéresse à des aspects plus généraux de cet élément génétique mobile qui pourraient se révéler très importants en terme de mouvement de gènes au sein des staphylocoques, voir d'autres bactéries gram-positives. Finalement ce travail de thèse met en place le fondamentaux requis pour des recherches futures visant à interférer avec le transfert horizontal de la résistance aux β-lactamines. - Staphylococcus aureus is a major human pathogen. Moreover, S. aureus have developed resistance to almost all available antibiotics, including the important family of β-lactam molecules. Intrinsic resistance to β-lactams is conferred by the Staphylococcal Cassette Chromosome mec (SCCmec), which is a mobile genomic island that inserts into the staphylococcal chromosome and can be horizontally transferred into other staphylococci. However, little is known about the molecular mechanisms involved in this horizontal transfer into naïve strains. One of the first steps in SCC mec horizontal transfer is its excision from the chromosome. Excision is mediated by recombinase enzymes that are encoded by SCC mec itself, and named accordingly Ccr recombinases - for Cassette Chromosome recombinases. One goal of this thesis was to understand the regulation these recombinase genes. By using original molecular tools we could demonstrate first that the Ccr recombinases were expressed in a "bistable" manner, i.e. in only few percentages of the bacterial cells at a given time, and second that they were regulated by determinants that were not encoded on the SCC mec element, but elsewhere on the staphylococcal genome. "Bistable" expression Ccr recombinases is an important concept. It allows SCC mec to be excised and thus available for horizontal transfer, while ensuring that only some cells, but not the whole population, loose their valuable SCC mec genes. Thus, while the SCC mec element expands with the multiplication of the MRSA colony, it can simultaneously be transmitted into methicillin-susceptible S. aureus (MSSA), which convert into new MRSA. Most interestingly, the fact that bistability was regulated by the cells, rather than by SCC mec, indicates that it was the choice of the bacteria to trigger or not SCC mec transfer. As a consequence, there must be, in nature, staphylococcal strains that are more or less prone to sustain SCC mec transfer. Following these seminal observations we found that excision was indeed tightly regulated during bacterial division, and occurred only during a limited period of time at the beginning of bacterial growth. This is compatible with cell-density mediated gene regulation, and may depend on the production of extracellular signal molecules that transmit appropriate orders to neighboring cells, such as in quorum sensing. The potential signal triggering SCCmec excision is as yet unknown. However, it could be critical in promoting the horizontal transfer of methicillin resistance, or for the possible development of means to interfere with it. Two additional hypothesis were logically investigated in the view of these first results. First, if some strains of MRSA might be more prone than others to promote SCC mec excision, then some strains of MS SA might be more or less prone to acquire the element as well. Second, to investigate these multiple mechanisms at an epidemiological level, one would need to develop tools amenable to explore S. aureus strains at a larger scale. Regarding the first issue, it was postulated by others that some MSSA might be refractory to SCC mec integration because they had peculiar DNA polymorphisms in the vicinity of the site-specific chromosomal entry point {attB) of SCC mec. By studying >40 S. aureus isolates from healthy carriers, we confirmed the polymorphism of the attB environment. Moreover, we could show that these polymorphic regions co-evolved with well-known phylogenic clonal clusters. Therefore, if SCCwec-refractory attB environments exist, then they would segregate in well- defined S. aureus clonal clusters that would be easy to identify at the epidemiological level. Regarding the second issue, we were able to construct a new excision reporter system in a low copy number S. aureus plasmid. The reporter system consists in a strong promoter driving a green fluorescent protein {gfp) gene, separated by an artificial SCC-like element carrying three transcriptional terminators. Thus, fluorescence is not expressed unless the SCC-like element is excised. The system has been successfully tested in several aureus and non- aureus staphylococci, and is now being applied to more strains and various excision- triggering or inhibiting conditions. Altogether the dissertation is a scientific journey through various aspects of a salient medical problem with regard to antibiotic resistance and public health threat. The research work tackles fundamental issues about the mechanisms of horizontal transfer of the SCC mec element. Moreover, it also addresses more general features of this mobile element, which could be of larger importance with regard to gene trafficking in staphylococci, and maybe other gram-positive bacteria. Finally, the dissertation sets the fundamentals for future work and possible new ways to interfere with the horizontal transfer of methicillin resistance.

Organization of lin genes and IS6100 among different strains of hexachlorocyclohexane-degrading Sphingomonas paucimobilis: evidence for horizontal gene transfer.

The organization of lin genes and IS6100 was studied in three strains of Sphingomonas paucimobilis (B90A, Sp+, and UT26) which degraded hexachlorocyclohexane (HCH) isomers but which had been isolated at different geographical locations. DNA-DNA hybridization data revealed that most of the lin genes in these strains were associated with IS6100, an insertion sequence classified in the IS6 family and initially found in Mycobacterium fortuitum. Eleven, six, and five copies of IS6100 were detected in B90A, Sp+, and UT26, respectively. IS6100 elements in B90A were sequenced from five, one, and one regions of the genomes of B90A, Sp+, and UT26, respectively, and were found to be identical. DNA-DNA hybridization and DNA sequencing of cosmid clones also revealed that S. paucimobilis B90A contains three and two copies of linX and linA, respectively, compared to only one copy of these genes in strains Sp+ and UT26. Although the copy number and the sequence of the remaining genes of the HCH degradative pathway (linB, linC, linD, and linE) were nearly the same in all strains, there were striking differences in the organization of the linA genes as a result of replacement of portions of DNA sequences by IS6100, which gave them a strange mosaic configuration. Spontaneous deletion of linD and linE from B90A and of linA from Sp+ occurred and was associated either with deletion of a copy of IS6100 or changes in IS6100 profiles. The evidence gathered in this study, coupled with the observation that the G+C contents of the linA genes are lower than that of the remaining DNA sequence of S. paucimobilis, strongly suggests that all these strains acquired the linA gene through horizontal gene transfer mediated by IS6100. The association of IS6100 with the rest of the lin genes further suggests that IS6100 played a role in shaping the current lin gene organization.

Is there a wage penalty for horizontal and vertical mismatch?

This paper studies how the horizontal and vertical mismatches in the labor market affect wage. We do so by taking into account that by choosing a job, wage and mismatches are simultaneously determined. The Seemingly Unrelated Equations model also allows us to control for any omitted variable that could cause biased estimators. We use REFLEX data for Spain. Results reveal that in most cases being horizontally matched has a wage premium and being over-educated does not affect wage. Results suggest that the modeling strategy successfully accounts for some omitted variable that affects simultaneously the probability of being horizontally matched and the wage. This could explain the existence of a wage penalty for over-educated workers when the omitted variable issue is not dealt with.

Horizontal innovation in the theory of growth and development

We analyze recent contributions to growth theory based on the model of expanding variety of Romer (1990). In the first part, we present different versions of the benchmark linear model with imperfect competition. These include the labequipment model, labor-for-intermediates and directed technical change . We review applications of the expanding variety framework to the analysis of international technology diffusion, trade, cross-country productivity differences, financial development and fluctuations. In many such applications, a key role is played by complementarities in the process of innovation.

Allocating ideas: Horizontal competition in tournaments

We develop a stylized model of horizontal and vertical competition intournaments with two competing firms. The sponsor cares about the qualityof the design but also about the design location. A priori not even thesponsor knows his preferred design location, which is only discoveredonce he has seen the actual proposals. We show that the more efficientfirm is more likely to be conservative when choosing the design location.Also, to get some differentiation in design locations, the cost differencebetween contestants can neither be too small nor too big. Therefore, ifthe sponsor mainly cares about the design location, participation in thetournaments by the two lowest cost contestants cannot be optimal for thesponsor.

Distribución horizontal de la anchoveta peruana con relación a variables ambientales en el período 1986 - 2000

Se analizó la distribución horizontal de la anchoveta Engraulis ringens utilizando sistemas de información geográfica. Los datos fueron obtenidos de los cruceros de evaluación hidroacústica de recursos pelágicos realizados durante los veranos de 1986 al 2000. Los resulitados indican que la distribución horizontal de la anchoveta está asociada a parámetros oceanográficos superficiales del mar, como rangos de temperatura, salinidad y clorofila a; sin embargo,parece ser que el parámetro más importante es la salinidad.

Horizontal and vertical distributions of zooplanckton numbers and biomass off the coast of Peru

Estudio de las distribuciones horizontales y verticales del zooplancton a lo largo de una línea desde cerca de la costa hasta el borde de la plataforma.

Distribución horizontal de los Amphipoda (crustacea) en el mar peruano durante el fenómeno "El niño" verano 1983

Los anfípodos hepéridos son microcrustáceos marinos que tienen un rol importante en la ecología trófica de especies pesqueras, se caracterizan por su voraz comportamiento predador sobre otros componentes del zooplancton (NELSON, 1979) y debido a su abundancia constituyen una fuente de alimento para niveles tróficos superiores (ALVAREZ y VIÑAS, 1994). El área de estudio comprende desde 03°29.80´S hasta los 12°54.50´S, abarcando los perfiles de Puerto Pizarro, Cabo Blanco, Talara, Paita, Punta Falsa, Pimentel, Chicama, Salaverry, Chimbote, Huarmey, Supe, Guacho, Chancay, Callao, Pucusana y Cerro Azul, cubriendo de esta manera la parte norte y centro del litoral peruano hasta las 100 millas náuticas de la costa y 50 metros de profundidad. Se determinan 34 especies de anfípodos, con 26 nuevos registros para el país y 2 especies nuevas. La familia Hyperridae es la que registra mayor representatividad a nivel específico destacando por su abundancia las especies Hyperioides sibaginis e Hyperietta vossleri. El presente trabajo, contribuye principalmente a ampliar el conocimiento de especies de anfípodos y asociarlas con las características ambientales, así las especies Vibilia chuni, Partaphronima gracilis, Hemithiphys tenuimanuss, Euscelus robustus y Schizoscelus ornatus, se encuentran relacionados con salinidades que fluctuan entre 34.8 y 35.1‰ y temperaturas de 25.4 hasta 28.2°C, rangos correspondientes a aguas de mezcla de Aguas Ecuatoriales Superficiales (AES) y Aguas Subtropicales superficiales (ASS).

Variabilidade horizontal de atributos de fertilidade e amostragem do solo no sistema plantio direto

As adubações, a lanço, quando desuniformes, ou as em linhas, no sistema plantio direto, aumentam a variabilidade dos atributos químicos do solo. Nestas condições, há necessidade de se saber qual a melhor forma e o número de subamostras por coletar para uma boa representatividade da fertilidade da área a ser cultivada. O objetivo deste estudo foi quantificar a variabilidade horizontal de atributos de fertilidade do solo no sistema plantio direto com diferentes modos de adubação e tempos de cultivo, com vistas em definir o número de subamostras necessárias para formar uma amostra representativa da fertilidade do solo de uma área. Foram coletadas, em novembro de 1997, 36 amostras simples, com pá de corte, na camada de 0-10 cm, de forma diferenciada nas adubações a lanço (5 e 10 cm, espessura da fatia e largura) e em linhas (5 cm de espessura da fatia pela largura das entrelinhas de semeadura da última cultura), em oito lavouras comerciais na região noroeste do Rio Grande do Sul. O número mínimo de subamostras para estimar com boa representatividade (α = 0,05 e erro em relação à média de 10%) o pH, o índice SMP e o teor de matéria orgânica foi baixo (menor do que 8) e alto (maior do que 40) para fósforo e potássio (Mehlich-1). O número de subamostras indicado para o sistema convencional (20) pode ser usado no sistema plantio direto, desde que seja admitido um erro de 20% em relação à média.

Variabilidade horizontal da fertilidade do solo de uma unidade de amostragem em sistema plantio direto

A coleta de amostras de solo com trado é mais simples e rápida do que com a pá de corte, sendo esta, freqüentemente, utilizada no sistema plantio direto para amostragem e determinação da fertilidade média do solo. Com o objetivo de comparar dois sistemas de amostragem localizada do solo (pá de corte ou trado de caneca) quanto à variabilidade horizontal de características químicas, determinar o número de unidades de amostra necessário à formação de uma amostra representativa da unidade de amostragem e demonstrar que a análise da amostra composta apresenta iguais resultados aos obtidos pelas médias aritméticas das análises das amostras simples, realizou-se um estudo em área há 15 anos sob sistema plantio direto. Os sistemas de amostragem localizada foram: pá de corte (24 amostras simples coletadas perpendicularmente ao sulco de plantio e no espaço compreendido entre os pontos médios entre sulcos) e trado (20 amostras simples coletadas no sulco de plantio (Ts), 20 amostras simples coletadas lateralmente a 10 cm do sulco (T10) e 20 amostras simples coletadas a 40 cm do sulco (T40)). Subamostras representativas das amostras simples foram misturadas, visando à formação de amostras compostas, tanto para pá de corte quanto para trado. Nas amostras simples e compostas foram determinados os teores de P, K, Ca2+, Mg2+ e Al3+ e os valores de pH, H + Al e P-rem. Foram calculadas as médias das determinações das amostras simples e das compostas, os desvios-padrão e os coeficientes de variação, sendo as médias comparadas pelo teste t de Student. Foi também calculado o número de amostras simples necessário à formação de uma amostra composta, para os dois sistemas de amostragem. Os resultados mostraram que apenas os teores de K e Mg2+ e os valores de H + Al diferiram de um método de amostragem para outro. Na amostragem com trado de caneca, a variabilidade de todas as características avaliadas foi maior do que na amostragem com pá de corte. A variabilidade foi baixa para pH, P-rem e H + Al e alta para P, Al3+, Mg2+, K e Ca2+. Dez amostras simples coletadas com pá de corte ou 27 amostras simples coletadas de forma localizada com trado de caneca (cinco amostras coletadas no sulco de plantio, nove coletadas a 10 cm do sulco e 13 coletadas no ponto médio entre sulcos) foram necessárias à formação de amostras compostas representativas. As análises das amostras compostas, tanto de pá de corte quanto de trado, apresentaram valores semelhantes às médias das análises das respectivas amostras simples, caracterizando corretamente a fertilidade média da unidade de amostragem.

Efficacy of the barbecue manoeuvre in benign paroxysmal vertigo of the horizontal canal

Five to ten percent of benign paroxysmal positional vertigo are caused by the horizontal semi-circular variant (h-BPPV). In this study, we reviewed the efficacy of the Barbecue repositioning manoeuvre in h-BPPV, and we assessed the possible effect of different factors on the outcome. Barbecue manoeuvre consists in turning the supine patient around his longitudinal axis toward the unaffected side until 360 degrees are accomplished. After every 90 degrees step the patient is maintained in the new position for 30 s. We reviewed 46 patients with h-BPPV, treated by barbecue rotation from 2003 to 2005. After the first Barbecue manoeuvre, the patients were followed-up at intervals of approximately 1 week and the rotation was repeated if h-BPPV persisted (up to three rotations). Factors assessed were age, gender, duration of symptoms before treatment and type of h-BPPV (canalolithiasis vs. cupulolithiasis). Fisher's exact test was used for the analysis. Results: 85% of patients (39/46) were cured after a maximum of 3 rotations. 74% (34/46) were cured after the first manoeuvre and 80% (37/46) after the second one. None of the evaluated factors did significantly affect the efficacy (P > 0.05). The Barbecue manoeuvre is an efficient treatment of h-BPPV demonstrating 85% cure rate after a maximum of three sessions. 74% of the patients are healed after one manoeuvre. The efficacy is not affected by the evaluated factors.

Espacialização vertical e horizontal dos indicadores de qualidade para um latossolo vermelho cultivado com citros

O objetivo deste trabalho foi verificar a forma de espacialização vertical e horizontal dos indicadores de qualidade química, física e hídrica de um Latossolo Vermelho distrófico textura média, cultivado com citros. O pomar de laranjeira Valência sobre limoeiro Cravo foi plantado em 1995, em Alto Paraná, região noroeste do Paraná. Em agosto de 2002, coletaram-se amostras deformadas e indeformadas de solo para realizar determinações químicas, físicas e hídricas. A amostragem foi feita em 24 transeções perpendiculares aos troncos das laranjeiras, em duas profundidades (0-0,20 e 0,20-0,40 m) e em três posições (projeção da copa da laranjeira, rodado da entrelinha e centro da entrelinha). Os indicadores avaliados foram submetidos às análises multivariadas, por meio dos componentes principais, e agrupados pelo método de Tocher. Os resultados demonstraram que a qualidade do solo apresentou espacialização heterogênea entre os indicadores de qualidade física, química e hídrica do solo, resultante das ações de manejo do pomar de laranjeira. A integração dos indicadores de qualidade física, química e hídrica do solo caracterizou a espacialização vertical da qualidade do solo (0-0,40 m) na projeção da copa da laranjeira e a espacialização vertical e horizontal (0-0,20 e 0,20-0,40 m) no rodado da entrelinha e centro da entrelinha. No rodado da entrelinha (0-0,20 m) ocorreu a maior restrição física e hídrica associada à melhor condição química do solo para as laranjeiras.

The interplay between relatedness and horizontal gene transfer drives the evolution of plasmid-carried public goods.

Plasmids carry a wide range of genes that are often involved in bacterial social behaviour. The question of why such genes are frequently mobile has received increasing attention. Here, we use an explicit population genetic approach to model the evolution of plasmid-borne bacterial public goods production. Our findings highlight the importance of both transmission and relatedness as factors driving the evolution of plasmid-borne public goods production. We partition the effects of plasmid transfer of social traits into those of infectivity and the effect of increased relatedness. Our results demonstrate that, owing to its effect on relatedness, plasmid mobility increases the invasion and stability of public goods, in a way not seen in individually beneficial traits. In addition, we show that plasmid transfer increases relatedness when public goods production is rare but this effect declines when production is common, with both scenarios leading to an increase in the frequency of plasmid-borne public goods. Plasmids remain important vectors for the spread of social genes involved in bacterial virulence thus an understanding of their dynamics is highly relevant from a public health perspective.