Transmission assumptions generate conflicting predictions in host-vector disease models: a case study in West Nile virus

This review synthesizes the conflicting outbreak predictions generated by different biological assumptions in host–vector disease models. It is motivated by the North American outbreak of West Nile virus, an emerging infectious disease that has prompted at least five dynamical modelling studies. Mathematical models have long proven successful in investigating the dynamics and control of infectious disease systems. The underlying assumptions in these epidemiological models determine their mathematical structure, and therefore influence their predictions. A crucial assumption is the host–vector interaction encapsulated in the disease-transmission term, and a key prediction is the basic reproduction number, R ₀. We connect these two model elements by demonstrating how the choice of transmission term qualitatively and quantitatively alters R ₀ and therefore alters predicted disease dynamics and control implications. Whereas some transmission terms predict that reducing the host population will reduce disease outbreaks, others predict that this will exacerbate infection risk. These conflicting predictions are reconciled by understanding that different transmission terms apply biologically only at certain population densities, outside which they can generate erroneous predictions. For West Nile virus, R ₀ estimates for six common North American bird species indicate that all would be effective outbreak hosts.     

Duelling timescales of host movement and disease recovery determine invasion of disease in structured populations

The epidemic potential of a disease is traditionally assessed using the basic reproductive number, R ₀. However, in populations with social or spatial structure a chronic disease is more likely to invade than an acute disease with the same R ₀, because it persists longer within each group and allows for more host movement between groups. Acute diseases 'perceive' a more structured host population, and it is more important to consider host population structure in analyses of these diseases. The probability of a pandemic does not arise independently from characteristics of either the host or disease, but rather from the interaction of host movement and disease recovery timescales. The R _* statistic, a group-level equivalent of R ₀, is a better indicator of disease invasion in structured populations than the individual-level R ₀.     

Temperature responses are a window to the physiology of dark respiration: differences between CO2 release and O2 reduction shed light on energy conservation

We showed that temperature responses of dark respiration for foliage of Pinus radiata could be approximated by Arrhenius kinetics, whereby E ₀ determines shape of the exponential response and denotes overall activation energy of respiratory metabolism. Reproducible and predictable deviation from strict Arrhenius kinetics depended on foliage age, and differed between R _CO2 and R _O2. Inhibition of oxygen reduction ( R _O2) by cyanide (inhibiting COX) or SHAM (inhibiting AOX) resulted in reproducible changes of the temperature sensitivity for R _O2, but did not affect R _CO2. Enthalpic growth – preservation of electrons in anabolic products – could be approximated with knowledge of four variables: activation energies ( E ₀) for both R _CO2 and R _O2, and basal rates of respiration at a low reference temperature ( R _REF). Rates of enthalpic growth by P. radiata needles were large in spring due to differences between R _REF of oxidative decarboxylation and that of oxygen reduction, while overall activation energies for the two processes were similar. Later during needle development, enthalpic growth was dependent on differences between E ₀ for R _CO2 as compared with R _O2, and increased E ₀( R _O2) indicated greater contributions of cytochrome oxidase to accompany the switch from carbohydrate sink to source. Temperature-dependent increments in stored energy can be calculated as the difference between R _CO2▵ H _CO2 and R _O2▵ H _O2.     

Flow cytometric DNA analysis of nurse shark, Ginglymostoma cirratum (Bonaterre) and clearnose skate, Raja eglanteria (Bosc) peripheral red blood cells

Flow cytometric DNA analysis was used to determine the DNA content of red blood cells from Ginglymostoma cirratum (Bonaterre) and Raja eglanteria (Bosc). The DNA content/nucleus was calculated to be 7.6 and 7.1 pg/nucleus, respectively. Peripheral red blood cells obtained both from G. cirratum and from R. eglanteria were demonstrated to be actively cell cycling. Percentages of red blood cells in the phases of the cell cycle ( G ₀/ G ₁, S , and G ₂/ M ) not only varied considerably between different fishes but also were quite variable in weekly samples obtained from individual animals.     

Non-association between Rfp-Y major histocompatibility complex-like genes and susceptibility to Marek's disease virus-induced tumours in 63× 72 F2 intercross chickens

Marek's disease (MD) is a lymphoproliferative disease caused by a member of the herpesvirus family, and the best understood genetic resistance to MD involves the chicken major histocompatibility complex (MHC) B -complex. Preliminary observations have suggested that MHC-like Rfp-Y genes might also influence the incidence of MD. This study describes the differentiation and definition of unique Rfp-Y genes in inbred lines 6₃ and 7₂, lines that possess identical B -complex genes, but that are resistant or susceptible to MD, respectively. To assess if Rfp-Y genes affect susceptibility to MD, 265 6₃× 7₂ F₂ chickens were challenged with the JM strain of MD virus at 1 week of age and were evaluated for MD lesions at up to 10 weeks of age. Genotyping of the F₂ chickens for Rfp-Y haplotypes was performed by restriction fragment length polymorphism analysis of genomic DNA using Taq I and a B-FIV probe. Analysis of variance and interval mapping procedures were used to determine association between the Rfp-Y haplotypes and the phenotypic MD values of the F₂ chickens. The cosegregation analysis of 265 F₂ chickens indicated that there was no association between Rfp-Y haplotypes and MD susceptibility. Furthermore, the fact that the Rfp-Y haplotypes fit the 1:2:1 segregation ratio and the Rfp-Y allele frequencies did not differ significantly from 0·5 in the full population or in selected subpopulations (of either 40 MD-resistant or 39 MD-susceptible chickens) also indicated that Rfp-Y haplotypes do not significantly influence MD susceptibility. We conclude that Rfp-Y haplotypes do not play a major role in determining the genetic susceptibility to MD in 6₃× 7₂ F₂ White Leghorn chickens.     

The importance of xylem constraints in the distribution of conifer species

Vulnerability of stem xylem to cavitation was measured in 10 species of conifers using high pressure air to induce xylem embolism. Mean values of air pressure required to induce a 50% loss in hydraulic conductivity (φ₅₀) varied enormously between species, ranging from a maximum of 14.2±0.6 MPa (corresponding to a xylem water potential of −14.2 MPa) in the semi-arid species Actinostrobus acuminatus to a minimum of 2.3±0.2 MPa in the rainforest species Dacrycarpus dacrydioides . Mean φ₅₀ was significantly correlated with the mean rainfall of the driest quarter within the distribution of each species. The value of φ₅₀ was also compared with leaf drought tolerance data for these species in order to determine whether xylem dysfunction during drought dictated drought response at the leaf level. Previous data describing the maximum depletion of internal CO₂ concentration (c_i) in the leaves of these species during artificial drought was strongly correlated with φ₅₀ suggesting a primary role of xylem in effecting leaf drought response. The possibility of a trade-off between xylem conductivity and xylem vulnerability was tested in a sub-sample of four species, but no evidence of an inverse relationship between φ₅₀ and either stem-area specific (K_a) or leaf-area specific conductivity (K₁) was found.     

Elasticity analysis in epidemiology: an application to tick-borne infections

The application of projection matrices in population biology to plant and animal populations has a parallel in infectious disease ecology when next-generation matrices (NGMs) are used to characterize growth in numbers of infected hosts ( R ₀). The NGM is appropriate for multi-host pathogens, where each matrix element represents the number of cases of one type of host arising from a single infected individual of another type. For projection matrices, calculations of the sensitivity and elasticity of the population growth rate to changes in the matrix elements has generated insight into plant and animal populations. These same perturbation analyses can be used for infectious disease systems. To illustrate this in detail we parameterized an NGM for seven tick-borne zoonoses and compared them in terms of the contributions to R ₀ from three different routes of transmission between ticks, and between ticks and vertebrate hosts. The definition of host type may be the species of the host or the route of infection, or, as was the case for the set of tick-borne pathogens, a combination of species and the life stage at infection. This freedom means that there is a broad range of disease systems and questions for which the methodology is appropriate.     

Responses of mosquitoes of the Anopheles farauti complex to 1-octen-3-ol and light in combination with carbon dioxide in northern Queensland, Australia

Abstract. In northern Queensland, Australia, three experiments were conducted to determine the response of mosquitoes of the Anopheles farauti complex to CDC traps baited with four attractant combinations: octenol + C0₂ and light; octenol and light; CO, and light; or C0₂ and octenol without light. A CDC-modified updraft light-trap was also trialled, but did not significantly enhance collections of An.farauti sensu lato. The combination of light, octenol and C0₂ caught significantly more An.farauti s.l. (both An.farauti No. 1 and No. 2 sibling species) when compared to C0₂ and light alone. Only small numbers of the An.farauti complex were captured when CDC traps were baited with octenol alone, i.e. no light or C0₂.     

Response properties of CO2-sensitive receptors in tsetse flies (Diptera: Glossina Palpalis)

Abstract. Extracellular single cell recordings of CO₂-sensitive receptors in tsetse flies revealed a steep dose response over a range of stimuli of two to three orders of magnitude and a maximum response of approximately 70 impulses/s after exposure to a high, but naturally feasible, C0₂ concentration of 5%. These receptor neurones are slightly sensitive to C0₂ levels occurring in air (-0.03%); the sensitivity to CO² above that level may be used to locate potential hosts. The C0₂-sensitive neurones did not respond to some other biologically relevant odours such as octenol, butanone or p-cresol; however, other receptor cells, some in the same sensillum, are sensitive to some of these odours.
A striking feature of the C0₂ receptors is that they appear not to adapt in their response frequencies. The spike numbers of the phasic-tonic response remain constant in the tonic portion during continuous (or repetitively pulsed) long-term stimulation (1 min). This unusual physiological ability would allow continuous monitoring of C0₂ values as well as detecting potential hosts that exhale CO₂. These electrophysiological results are compared to the behavioural findings in tsetse flies and to the physiological data on C0₂ receptors of other, non-bloodsucking insects.     

A microsatellite analysis of natterjack toad, Bufo calamita, metapopulations

Although it is widely recognised that spatial subdivision of populations is common in nature, there is no consensus as to how metapopulation dynamics affect genetic diversity. We investigated the genetic differentiation of natterjack toads, Bufo calamita , in three regions of Britain where habitat continuity indicated the likely occurrence of extensive metapopulations. Our intention was to determine whether genetic analysis supported the existence of metapopulation structures, if so of what type, and to identify barriers to migration between subpopulations. Allele frequencies were determined across eight polymorphic microsatellite loci for a total of 24 toad subpopulations at three separate sites. Genetic differentiation was assessed using five measures of genetic distance, notably F _ST , R _ST , Nei's standard distance D _s , Δμ² and the Cavalli-Sforza chord distance D _c . B. calamita exhibited small but significant levels of genetic differentiation between subpopulations in all three study areas, and genetic and geographic distance correlations indicated isolation-by-distance effects in all three cases. The effects on correlation strengths of compensation for positive (sea, rivers, urban development) and negative (pond clusters) barriers to toad migration between the subpopulations in each area were also determined. D _c , a measure which assumes that differentiation is caused by drift with negligible mutation effect, yielded the most plausible interpretation of metapopulation structures. Overall the patterns of genetic variation suggested the existence of a mixed metapopulation model for this species, with high levels of gene flow compatible with one version of the classical model but often supported by particularly stable subpopulations as in the mainland-island model.     

Vector-borne diseases and the basic reproduction number: a case study of African horse sickness

Abstract. The basic reproduction number, R ₀, can be used to determine factors important in the ability of a disease to invade or persist. We show how this number can be derived or estimated for vector-borne diseases with different complicating factors. African horse sickness is a viral disease transmitted mainly by the midge Culicoides imicola. We use this as an example of such a vector-transmitted disease where latent periods, seasonality in vector populations, and multiple host types may be important. The effect of vector population dynamics which are dependent on either host or vector density are also addressed. If density-dependent constraints on vector population density are less severe, R_o is more sensitive to vector mortality and the virus development rate. Host-dependent vector dynamics change the relationship between R₀ and host population size. Seasonality can either increase or decrease the estimate of R₀ , depending on the lag between the peak of the midge population and the infective host population. The relative abundance of two host types is a factor in the ability of a disease to invade, but the strength of this factor depends on the differences between the hosts in recovery from infection, mortality and transmission. Removal of a reservoir host may increase R_Q.     

LEAF INFECTION OF COTTON BY XANTHOMONAS MALVACEARUM (E.F.SM.) DOWSON

Xanthomonas malvacearum spread more rapidly along vascular tissue than into mesophyll when inoculated to the main veins of susceptible cotton leaves. The extent of spread varied with the concentrations of inocula, tissue age and cotton variety.
Increasing concentrations of inocula accelerated the initial spread of disease.
Bacteria spread more rapidly in young leaves than in old—increasing age greatly decreased disease in the mesophyll. The initial invasion was quicker in young leaves of young plants than in young leaves of old plants.
Three types of behaviour, according to the host's reaction, distinguish Knight's resistance factors: ( a ) where X. malvacearum spread extensively along veins and into mesophyll of plants containing factors B₃ and B₅; ( b ) where it was restricted to the point of inoculation in plants containing B ₄, B₉ and combinations with B _6m; and ( c ) where it spread along veins but not appreciably into mesophyll in varieties containing B ₂ and B ₂ B ₃.
From this and four other different types of tests, factors B ₂ and B ₃ seem to increase mesophyll resistance but only B ₂ gives appreciable vascular resistance. Further, the vascular bundles in varieties with B ₂ seem to be surrounded by an additional 'barrier' which resists X. malvacearum.     

A Chemical Indicator for the Rapid Measurement of F0 Values

A new chemical indicator for monitoring steam sterilization processes has been calibrated in F ₀ units. The effective temperature range for F ₀ measurements using this device has been shown to lay between 115 and 123°C. The effective F ₀ range of the device has been shown to be 4–23 F ₀ units. Using the device, measurements can be made within 0.5 units of conventionally calculated F ₀ values.     

Age, growth and reproduction of the barrelfish Hyperoglyphe perciformis (Mitchill) in the western North Atlantic

Otoliths ( n = 847) and gonads ( n = 817) were collected from barrelfish Hyperoglyphe perciformis that were captured by commercial fishermen in the waters off South Carolina and Georgia in 1995, 1997 and 2001–2006. Of the otoliths collected, 97% were aged successfully, and specimens sampled ranged from 5 to 85 years, with a median age of 12 years. The von Bertalanffy growth parameters yielded the equation: L_t = 857·8{1 − e^{−0·0985[ t −(−8·95)]}}, where L_t is fork length ( L _F) at time t . Through histological examination, 94% of the gonads assessed were assigned to a sex and reproductive class. Females spawned from September to May with a peak from November to January. Males spawned year round, but had a peak from September to April. The sex ratio (M:F) for this population was 1:1·34. The smallest mature female was 605 mm L _F and the youngest immature female was 697 mm L _F. Estimates of L _F and age at 50% maturity ( L ₅₀ and A ₅₀) for females were 660 mm L _F (95% CI = 633–667 mm L _F) and 6·08 years (95% CI = 3·50–7·27 years), respectively. The youngest mature male was 575 mm L _F and the oldest immature male was 762 mm L _F, and no estimates of L ₅₀ or A ₅₀ were made for males. It was determined that barrelfish exhibit the typical characteristics of long life span, slow growth and high age at maturity seen in other deepwater fishes, and that care should be taken to manage this species accordingly.     

Three parameters comprehensively describe the temperature response of respiratory oxygen reduction

Using an exponential model that relies on Arrhenius kinetics, we explored Type I, Type II and dynamic (e.g. declining Q ₁₀ with increasing temperature) responses of respiration to temperature. Our Arrhenius model provides three parameters: R _REF (the base of the exponential model, nmol g⁻¹ s⁻¹), E ₀ (the overall activation energy of oxygen reduction that dominates its temperature sensitivity, kJ mol⁻¹) and δ (that describes dynamic responses of E ₀ to measurement temperature, 10³ K²). Two parameters, E ₀ and δ , are tightly linked. Increases in overall activation energy at a reference temperature were inversely related to changes in δ . At an E ₀ of ca. 45 kJ mol⁻¹, δ approached zero, and respiratory temperature response was strictly Arrhenius-like. Physiologically, these observations suggest that as contributions of AOX to combined oxygen reduction increase, E ₀( REF ) decreases because of different temperature sensitivities for V _max, and δ increases because of different temperature sensitivities for K _1/2 of AOX and COX. The balance between COX and AOX activity helps regulate plant metabolism by adjusting the demand for ATP to that for reducing power and carbon skeleton intermediates. Our approach enables determination of respiratory capacity in vivo and opens a path to development of process-based models of plant respiration.     

Ecophysiological and morphological parameters related to survival in grass species exposed to an extreme climatic event

An experiment was performed to elucidate interspecific differences in survival time of grass species subjected to an extreme climatic event. We exposed eight grass species to a simulated heat wave in the field ('free air' temperature increase at 11°C above ambient) combined with drought. We determined whether interspecific differences in survival time were related to the responses of the species to the imposed stress or could be explained by their ecophysiological or morphological characteristics in unstressed conditions. Surprisingly, there was no effect of specific leaf area, but species with a higher total leaf area survived longer. This may arise from a greater water reserve in the plant as a whole, which could delay the desiccation of the meristem, or from reduced evaporation due to a higher leaf area index. Species in which the decrease in light-saturated stomatal conductance ( g _s ) and photosynthetic CO₂ uptake rate ( A _max ) was strongly related to the decrease in soil water availability (measured as soil relative water content and stress duration) survived longer than species in which g _s and A _max likewise declined but responded more to daily fluctuations in irradiance, temperature, and vapor pressure deficit during the heat wave. We, therefore, hypothesize that interspecific differences in stress survival time might be related to the extent to which stomata react to changes in soil water conditions relatively to changes in other environmental and physiological factors. The results suggest that resistance to extremes is governed by other mechanisms than resistance to moderate drought.     

Age and growth analysis of the central mudminnow, Umbra limi (Kirtland, 1840)

A rudimentary understanding of age, growth, and life-span is lacking for many non-game fishes. Growth characteristics of the central mudminnow ( Umbra limi ) have not yet been accurately described using reliable hard part analysis. The utility of scales and otoliths as ageing structures and quantified growth was examined in one lake and one stream population of central mudminnow. Scales were found to be of no utility in determining age due to inconsistent formation of yearly annuli and a high incidence of regenerated scales, while otoliths were easily extracted and considered to be an accurate ageing structure. Ages determined from scales were low compared to those from otoliths, and the difference in age interpreted from the two structures increased with fish age. A power function was fitted to describe the length-weight relationship for this species ( a  = 0.0069, b  = 3.175). Von Bertalanffy growth parameters were estimated and compared for each population (Lake: L _∞ = 114.20 mm, K  = 0.30, t ₀ = −0.93; Stream: L _∞ = 77.59 mm, K  = 0.63, t ₀ = −0.76). The lake population showed greater size at age compared to the stream population, especially at older ages, and achieved a larger maximum size. Growth rate was also greater in the lake population (Lake: 1.74; Stream: 1.09 g year⁻¹). Females were larger at age than males in both populations, however all individuals greater than age 3 were males. This work represents the first successful account of central mudminnow growth using hard part analysis.     

Occurrence of aminopropylcadaverine and its aminopropyl derivatives aminopentylnorspermidine and N, N' -bis(3-aminopropyl)cadaverine in Halococcus acetoinfaciens

Abstract Besides putrescine, cadaverine, spermidine, spermine and thermospermine, three novel polyamines were detected in a slightly halophilic eubacterium Halococcus acetoinfaciens (IAM 12094, ATCC 25861). These novel polyamines were found to be N -3-aminopropylcadaverine [NH₂(CH₂)₃NH(CH₂)₅NH₂] and its aminopropyl derivatives: aminopentylnorspermidine [NH₂(CH₂)₃NH(CH₂)₃NH(CH₂)₅NH₂] and N , N ' -bis(3-aminoprophyl)cadaverine [NH₂(CH₂)₃ NH(CH₂)₅NH(CH₂)₃NH₂]. Aminopropylcadaverine was also detected in two other species, Halococcus agglomeratus (IAM 12095, ATCC 25862) and Halococcus nondenitrificans (IAM 12096, ATCC 25863).