RESUMO
Many myodocopid ostracods are unusual in that they have well-developed compound eyes yet must view their environment through a shell. The cypridinid Macrocypridina castanea is relatively large among ostracods (about 5-10 mm) and is a pelagic predator. This species possess highly pigmented shells with a transparent region lying just above the eye. Here we examine the ultrastructure and transparency of this window using electron microscopy, serial-block face scanning electron microscopy and X-ray diffraction analysis and optical modelling. An internal, laminar stack was identified within the window region of the shell that formed a more regular half-wave reflector than in non-window regions, and where the distance between molecules in the chitin-protein fibrils decreases as compared to the non-window area. This results in excellent transmission properties-at around 99% transmission-for wavelengths between 350 and 630 nm due to its half-wave reflector organization. Therefore, blue light, common in the mid and deep sea, where this species inhabits, would be near-optimally transmitted as a consequence of the sub-micrometre structuring of the shell, thus optimizing the ostracod's vision. Further, pore canals were identified in the shell that may secrete substances to prevent microbial growth, and subsequently maintain transparency, on the shell surface. This article is part of the theme issue 'Bioinspired materials and surfaces for green science and technology'.
Assuntos
Crustáceos/anatomia & histologia , Olho , Fenômenos Ópticos , Exoesqueleto/anatomia & histologia , Animais , FemininoRESUMO
Vaccine-specific antibody responses are essential in the diagnosis of antibody deficiencies. Responses to Pneumovax II are used to assess the response to polysaccharide antigens, but interpretation may be complicated. Typhim Vi® , a polysaccharide vaccine for Salmonella typhoid fever, may be an additional option for assessing humoral responses in patients suspected of having an immunodeficiency. Here we report a UK multi-centre study describing the analytical and clinical performance of a Typhi Vi immunoglobulin (Ig)G enzyme-linked immunosorbent assay (ELISA) calibrated to an affinity-purified Typhi Vi IgG preparation. Intra- and interassay imprecision was low and the assay was linear, between 7·4 and 574 U/ml (slope = 0·99-1·00; R2 > 0·99); 71% of blood donors had undetectable Typhi Vi IgG antibody concentrations. Of those with antibody concentrations > 7·4 U/ml, the concentration range was 7·7-167 U/ml. In antibody-deficient patients receiving antibody replacement therapy the median Typhi Vi IgG antibody concentrations were < 25 U/ml. In vaccinated normal healthy volunteers, the median concentration post-vaccination was 107 U/ml (range 31-542 U/ml). Eight of eight patients (100%) had post-vaccination concentration increases of at least threefold and six of eight (75%) of at least 10-fold. In an antibody-deficient population (n = 23), only 30% had post-vaccination concentration increases of at least threefold and 10% of at least 10-fold. The antibody responses to Pneumovax II and Typhim Vi® correlated. We conclude that IgG responses to Typhim Vi® vaccination can be measured using the VaccZyme Salmonella typhi Vi IgG ELISA, and that measurement of these antibodies maybe a useful additional test to accompany Pneumovax II responses for the assessment of antibody deficiencies.
Assuntos
Imunidade Adaptativa/imunologia , Anticorpos Antibacterianos/sangue , Ensaio de Imunoadsorção Enzimática/métodos , Imunoglobulina G/sangue , Síndromes de Imunodeficiência/diagnóstico , Polissacarídeos Bacterianos/imunologia , Vacinas Tíficas-Paratíficas/imunologia , Adolescente , Adulto , Idoso , Idoso de 80 Anos ou mais , Anticorpos Antibacterianos/imunologia , Formação de Anticorpos/imunologia , Feminino , Humanos , Imunoglobulina G/imunologia , Síndromes de Imunodeficiência/imunologia , Masculino , Pessoa de Meia-Idade , Vacinas Pneumocócicas/imunologia , Salmonella typhi/imunologia , Vacinação , Adulto JovemRESUMO
An extended spectroscopic study on the left-through-left circularly polarized reflection spectra of a large number of beetles from the Australasian Scrabaeidae:Cetoniinae of the Lomaptera genus was undertaken. We have obtained a five-category spectral classification. The principal spectral features, which even within the genus range from blue to infrared, are related to structural chirality in the beetle shells. The detailed features of each spectral classification are related to different structural perturbations of the helix, including various pitch values and abrupt twist defects. These spectral characteristics and associated shell structures are confirmed on the basis of simple modelling. An important conclusion from our study is that the simple helical structure resulting in a single symmetric Bragg peak is not the dominant spectral type. Rather the reality is a rich tapestry of spectral types. One intriguing specimen is identified via a scanning electron micrograph to consist of a double interstitial helix leading to a particular double-peak spectrum.
Assuntos
Estruturas Animais , Besouros , Luz , AnimaisRESUMO
Three species of cactus whose spines act as dew harvesters were chosen for this study: Copiapoa cinerea var. haseltoniana, Mammillaria columbiana subsp. yucatanensis and Parodia mammulosa and compared with Ferocactus wislizenii whose spines do not perform as dew harvesters. Time-lapse snapshots of C. cinerea showed movement of dew droplets from spine tips to their base, even against gravity. Spines emanating from one of the areoles of C. cinerea were submerged in water laced with fluorescent nanoparticles and this particular areole with its spines and a small area of stem was removed and imaged. These images clearly showed that fluorescent water had moved into the stem of the plant. Lines of vascular bundles radiating inwards from the surface areoles (from where the spines emanate) to the core of the stem were detected using magnetic resonance imaging, with the exception of F. wislizenii that does not harvest dew on its spines. Spine microstructures were examined using SEM images and surface roughness measurements (Ra and Rz) taken of the spines of C. cinerea It was found that a roughness gradient created by tapered microgrooves existed that could potentially direct surface water from a spine tip to its base.This article is part of the themed issue 'Bioinspired hierarchically structured surfaces for green science'.
Assuntos
Cactaceae/anatomia & histologia , Cactaceae/química , Estruturas Vegetais/anatomia & histologia , Estruturas Vegetais/química , Água/química , Anisotropia , Difusão , GravitaçãoAssuntos
Anticorpos Antibacterianos/imunologia , Formação de Anticorpos/imunologia , Síndromes de Imunodeficiência/imunologia , Polissacarídeos Bacterianos/imunologia , Salmonella typhi/imunologia , Vacinas Tíficas-Paratíficas/imunologia , Adulto , Agamaglobulinemia/imunologia , Idoso , Imunodeficiência de Variável Comum/imunologia , Ensaio de Imunoadsorção Enzimática , Feminino , Interações Hospedeiro-Patógeno/efeitos dos fármacos , Interações Hospedeiro-Patógeno/imunologia , Humanos , Imunoglobulina G/imunologia , Masculino , Pessoa de Meia-Idade , Estudos Prospectivos , Salmonella typhi/fisiologia , Febre Tifoide/imunologia , Febre Tifoide/microbiologia , Febre Tifoide/prevenção & controle , Vacinas Tíficas-Paratíficas/administração & dosagem , Vacinação/métodos , Adulto JovemRESUMO
Four species of cacti were chosen for this study: Copiapoa cinerea var. haseltoniana, Ferocactus wislizenii, Mammillaria columbiana subsp. yucatanensis and Parodia mammulosa. It has been reported that dew condenses on the spines of C. cinerea and that it does not on the spines of F. wislizenii, and our preliminary observations of M. columbiana and P. mammulosa revealed a potential for collecting dew water. This study found all four cacti to harvest dew on their stems and spines (albeit rarely on the spines of F. wislizenii). Dew harvesting experiments were carried out in the UK, recording an increase in cacti mass on dewy nights. By applying a ranking relative to a polymethyl methacrylate (Plexiglas) reference plate located nearby, it was found that C. cinerea collected the most airborne moisture followed by M. columbiana, P. mammulosa and F. wislizenii respectively, with mean efficiency ratio with respect to the Plexiglas reference of 3.48 ± 0.5, 2.44 ± 0.06, 1.81 ± 0.14 and 1.27 ± 0.49 on observed dewy nights. A maximum yield of normalized performance of 0.72 ± 0.006 l/m(-2) on one dewy night was recorded for C. cinerea. Removing the spines from M. columbiana was found to significantly decrease its dew harvesting efficiency. The spines of three of the species were found to be hydrophilic in nature, while F. wislizenii was hydrophobic; the stems of all four species were hydrophilic. The results of this study could be translated into designing a biomimetic water collecting device that utilizes cactus spines and their microstructures.
Assuntos
Materiais Biomiméticos/química , Cactaceae/química , Cactaceae/classificação , Caules de Planta/química , Caules de Planta/classificação , Água/química , Adsorção , Teste de Materiais , Especificidade da EspécieRESUMO
Some tenebrionind beetles inhabiting the Namib desert are known for using their body to collect water droplets from wind-blown fogs. We aim to determine whether dew water collection is also possible for desert insects. For this purpose, we investigated the infra-red emissivity, and the wetting and structural properties, of the surface of the elytra of a preserved specimen of Physasterna cribripes (Tenebrionidæ) beetle, where the macro-structure appears as a series of "bumps", with "valleys" between them. Dew formation experiments were carried out in a condensation chamber. The surface properties (infra-red emissivity, wetting properties) were dominated by the wax at the elytra surface and, to a lower extent, its micro-structure. We performed scanning electron microscope on histological sections and determined the infra-red emissivity using a scanning pyrometer. The emissivity measured (0.95±0.07 between 8-14 µm) was close to the black body value. Dew formation occurred on the insect's elytra, which can be explained by these surface properties. From the surface coverage of the condensed drops it was found that dew forms primarily in the valleys between the bumps. The difference in droplet nucleation rate between bumps and valleys can be attributed to the hexagonal microstructure on the surface of the valleys, whereas the surface of the bumps is smooth. The drops can slide when they reach a critical size, and be collected at the insect's mouth.
Assuntos
Besouros/fisiologia , Pele/ultraestrutura , Água/química , Exoesqueleto/química , Exoesqueleto/efeitos da radiação , Exoesqueleto/ultraestrutura , Animais , Besouros/ultraestrutura , Clima Desértico , Raios Infravermelhos , Modelos Teóricos , Transição de Fase , Pele/química , Pele/efeitos da radiação , Temperatura , Ceras/análise , Tempo (Meteorologia) , MolhabilidadeRESUMO
Nature has adapted different methods for surviving dry, arid, xeric conditions. It is the focus of this comparative review to pull together the relevant information gleaned from the literature that could be utilized to design moisture harvesting devices informed by biomimetics. Most water harvesting devices in current use are not informed by nature and those that do are usually based on a biomimetic principle that has been based on one species only. This review draws on the published literature to establish a list of species (animals (vertebrates/invertebrates) and plants) whose habitat is in mainly dry or arid regions and that are known to harvest airborne moisture. Key findings have been outlined and review comments and discussion set out. Following this, surface feature convergences have been identified, namely hexagonal microstructures, groove-like and cone-like geometries. This has been coupled with direction of water flow that is driven by surface energy. As far as the authors are aware, this convergent evolution has not been brought together in this manner before. In the future this information could be translated into an engineered device for collecting water from airborne sources.
Assuntos
Plantas/química , Plantas/metabolismo , Absorção Cutânea/fisiologia , Pele/química , Pele/metabolismo , Água/química , Água/metabolismo , Animais , Difusão , Humanos , Modelos Biológicos , Modelos Químicos , Propriedades de SuperfícieRESUMO
BNP7787, an investigational drug undergoing global Phase III development, appears to have potential advantages over other cytoprotective compounds that have been evaluated for preventing and mitigating cisplatin-induced nephrotoxicity. Herein, we characterized the in vitro accumulation of BNP7787 in human renal proximal tubule cells (HK-2) in which cisplatin is known to be taken up and accumulate. HK-2 cells were incubated with pharmacological concentrations of BNP7787 for varying times. Temperature-dependent accumulation of BNP7787 in cells was observed and the BNP7787-derived metabolite, mesna, formed intracellularly was directly monitored. The peak level of BNP7787-derived mesna measured in HK-2 cells was approximately 0.6 nmol/10(6) cells; this is pharmacologically similar to reported platinum concentrations in kidney cells and may be sufficient to afford nephroprotection. Therefore, in addition to previously suggested glomerular filtration, the cellular accumulation of BNP7787 by HK-2 cells is a plausible newly identified mechanism by which BNP7787 may accumulate in renal tubular cells, where it can exert its pharmacological effects to protect against cisplatin-induced nephrotoxicity by direct covalent conjugation of mesna with cisplatin, or by the formation of BNP7787-derived mesna-disulfide heteroconjugates that exert nephroprotective effects by inhibition of the key toxification enzyme targets γ-glutamyltranspeptidase and aminopeptidase N.
Assuntos
Túbulos Renais Proximais/metabolismo , Mesna/análogos & derivados , Linhagem Celular , Cromatografia Líquida de Alta Pressão , Eletroquímica , Humanos , Túbulos Renais Proximais/citologia , Mesna/farmacocinéticaRESUMO
PURPOSE: Previous studies from our laboratory have identified a role for gamma-glutamyl transpeptidase (GGT) in BNP7787 (disodium 2,2'-dithio-bis ethane sulfonate, dimesna, Tavocept™)-mediated cisplatin nephroprotection. Dekant has proposed that gamma-glutamyl transpeptidase (GGT), aminopeptidase N (APN) and cysteine-conjugate-ß-lyase (CCBL) comprise a multi-enzyme pathway that acts on xenobiotic-glutathione conjugates converting them to nephrotoxic metabolites. We report modulation of APN activity within this pathway by BNP7787-derived mesna-disulfide heteroconjugates. METHODS: A fluorimetric assay was used to determine the effect of BNP7787, BNP7787-derived mesna-disulfide heteroconjugates, and the BNP7787 metabolite, mesna (sodium 2-mercaptoethane sulfonate), on the initial velocity and overall progress curve of the human APN reaction in vitro. RESULTS: Neither BNP7787 nor mesna-cysteinyl-glutamate inhibited human APN. Select BNP7787-derived mesna-disulfide heteroconjugates (mesna-cysteine, mesna-glutathione, mesna-cysteinyl-glycine) and high concentrations of mesna inhibited APN activity. Allosteric effects on the enzyme progress curve outside of the linear initial velocity region were observed for mesna-cysteinyl-glycine, mesna-glutathione and mesna-cysteinyl-glutamate and appeared to be a function of having both mesna and di- or tri-peptide functionalities in one molecule. In situ-generated mesna-cisplatin conjugates were not a substrate for human APN. CONCLUSIONS: BNP7787-mediated prevention or mitigation of cisplatin-induced nephrotoxicity may involve APN inhibition by certain BNP7787-derived mesna-disulfide heteroconjugates and appears correlated to the presence of a glycinate moiety and/or an anionic group. Two general mechanisms for BNP7787-mediated nephroprotection of cisplatin-induced nephrotoxicity involving the GGT, APN and CCBL nephrotoxigenic pathway are proposed which acting in a concerted and/or synergistic manner, and thereby prevent or mitigate cisplatin-induced renal toxicity.
Assuntos
Antígenos CD13/antagonistas & inibidores , Cisplatino/toxicidade , Nefropatias/induzido quimicamente , Nefropatias/prevenção & controle , Mesna/análogos & derivados , Substâncias Protetoras/farmacologia , Regulação Alostérica , Biocatálise/efeitos dos fármacos , Antígenos CD13/genética , Antígenos CD13/metabolismo , Cisplatino/análogos & derivados , Cisplatino/metabolismo , Cisplatino/farmacologia , Cisteína/análogos & derivados , Cisteína/metabolismo , Cisteína/farmacologia , Dipeptídeos/farmacologia , Glutationa/análogos & derivados , Glutationa/metabolismo , Glutationa/farmacologia , Glicina/análogos & derivados , Glicina/metabolismo , Glicina/farmacologia , Humanos , Cinética , Mesna/metabolismo , Mesna/farmacologia , Modelos Biológicos , Substâncias Protetoras/metabolismo , Proteínas Recombinantes/antagonistas & inibidores , Proteínas Recombinantes/genética , Proteínas Recombinantes/metabolismoRESUMO
The structurally coloured chelicerae of jumping spiders (Salticidae) display some of the most striking of all colours in this family, in which they predominantly occur. Remarkably, however, the source of this iridescence has not been studied. For this reason, we chose to investigate the green iridescent chelicerae of the red-backed jumping spider, Phidippus johnsoni. The colour is restricted to the dorsal region of the basal chelicera segment--the paturon. This was confirmed by reflectance measurements taken at normal incidence and in backscatter, which gave a peak reflectance in the green (520 nm), arising from the first harmonic of a Bragg resonance in the near infrared. Transmission electron microscope analysis of the paturon cuticle revealed a stack of 86 layers of alternating low and high density materials, identified as air and chitin respectively. Simulations based on a periodic multilayer model of the ten outermost layers of this structure gave theoretical reflectance spectra, closely matching those observed, suggesting that the stack functions as a multilayer reflector for green. The colour is thought to function as a conspecific signal, since studies of vision in a closely related species, also displaying green chelicerae, have shown that the eyes have a peak spectral sensitivity, matching that of the chelicerae.
Assuntos
Aranhas/anatomia & histologia , Comunicação Animal , Animais , Cor , Microscopia Eletrônica de Varredura , Microscopia Eletrônica de Transmissão , Modelos Teóricos , Análise Espectral , Aranhas/fisiologia , Aranhas/ultraestruturaRESUMO
The technical and diagnostic performances of five commercially available enzyme-linked immunosorbent assays for the measurement of anti-diphtheria toxoid IgG antibodies were evaluated. There was good agreement between the relative sensitivities of the five assays, but the relative specificity of one of the assays differed from that of the other four assays. Three of the five assays possessed recoveries of the international reference material NIBSC 00/496 within the range of 90% to 110% at antibody levels >0.1 IU/ml. The data suggest that there are manufacture-dependent differences in relative sensitivity, specificity, and accuracy for the determination of anti-diphtheria toxoid IgG antibodies that could result in different diagnostic interpretations.
Assuntos
Anticorpos Antibacterianos/sangue , Antitoxinas/sangue , Técnicas Bacteriológicas/métodos , Toxoide Diftérico , Difteria/diagnóstico , Imunoglobulina G/sangue , Ensaio de Imunoadsorção Enzimática/métodos , Humanos , Sensibilidade e EspecificidadeRESUMO
Five commercially available enzyme-linked immunosorbent assays for the measurement of anti-tetanus toxoid immunoglobulin G (IgG) antibodies were evaluated for performance. The data suggest that there are manufacturer-dependent differences in sensitivity and accuracy for the determination of tetanus toxoid IgG antibodies that could result in different diagnostic interpretations.
Assuntos
Imunoglobulina G/sangue , Antitoxina Tetânica/sangue , Toxoide Tetânico/imunologia , Tétano/imunologia , Anticorpos Antibacterianos/sangue , Ensaio de Imunoadsorção Enzimática , Guias como Assunto , Humanos , Kit de Reagentes para Diagnóstico , Padrões de Referência , Sensibilidade e Especificidade , Tétano/microbiologiaRESUMO
The photonic structures of butterfly wings are among the most anatomically diverse of all those in nature, giving rise to an unrivalled display of structural colours. These have recently become the focus of research by workers in a variety of disciplines, stimulated by their potential applications to technology ('biomimetics'). This interest, together with the discovery of unpublished electron micrographs taken by the late Dr John Huxley (Natural History Museum, London), prompted this review of butterfly photonics in general. The current work provides a synopsis of the literature to date, covering the diversity and evolution of these optical structures and incorporating Huxley's work, which represents an important biomimetic and evolutionary database on its own. This review deals with butterfly photonic devices according to the parts of the butterfly scales on which they occur. In this way, the information is ripe for evolutionary study.
Assuntos
Biodiversidade , Evolução Biológica , Borboletas/genética , Borboletas/ultraestrutura , Luz , Animais , Asas de Animais/ultraestruturaRESUMO
An early step in the carcinogenesis of hereditary non-polyposis colorectal cancer (HNPCC) and some sporadic colorectal cancers (CRCs) is the acquisition of a 'mutator phenotype' resulting from defects in DNA mismatch repair (MMR) genes, which normally maintain genomic stability. This mutator phenotype causes an approximately 100-1000-fold increase in base substitutions and small insertion/deletion mutations thereby driving carcinogenesis. It also causes genome-wide microsatellite instability (MSI) due to the inability to repair mutations within these small, hard to replicate, repetitive DNA elements. In contrast, less is known about the role of mutator phenotypes in microsatellite stable (MSS) CRC. In this report, we have measured the mutation rates in 11 MSS CRC cell lines to obtain an estimate of the prevalence of mutator phenotypes in MSS carcinogenesis. Of the 11 cell lines, three of them (27%) possess spontaneous hypoxanthine phosphoribosyltransferase mutation rates approximately 10-100-fold above background. When challenged with alkylating and oxidising agents, the degree of survival and apoptotic responses are different, indicating that these cell lines may represent more than one mutator phenotype. These data demonstrate that a significant portion of MSS CRC cell lines has increased mutation rates and that this may play a role in MSS CRC carcinogenesis.
Assuntos
Apoptose , Transformação Celular Neoplásica , Neoplasias Colorretais/genética , Neoplasias Colorretais/patologia , Análise Mutacional de DNA , Repetições de Microssatélites , Alquilantes/farmacologia , Sobrevivência Celular , Reparo do DNA , Humanos , Oxirredução , Fenótipo , Células Tumorais CultivadasRESUMO
Faithful maintenance of the genome is crucial to the individual and the species. Oxidative DNA damage, such as 8-oxo-7,8-dihydroguanine (8-oxoG), poses a major threat to genomic integrity. 8-OxoG can mispair with 2'-deoxycytidine 5'-triphosphate or with 2'-deoxyadenosine triphosphate during DNA replication, forming C*8-oxoG and A*8-oxoG mispairs. Human MutY is responsible for recognition and removal of the inappropriately inserted adenine in an A*8-oxoG mispair. If unrepaired, the A*8-oxoG mispairs can result in deleterious C:G to A:T transversions. Human MutY functions in a postreplication repair pathway and is targeted to the newly synthesized daughter strand of DNA for removal of the adenine base. The human MutY protein is targeted to both the mitochondria and the nucleus and associates with the proliferating cell nuclear antigen, apurinic/ apyrimidinic endonuclease 1, replication protein A and mutS homolog 6 proteins. Mutations in the human MutY gene and defective activity of the human MutY protein have been detected in cancer. A direct correlation between defective A*8-oxoG repair and increased levels of genomic 8-oxoG has now been established.
Assuntos
DNA Glicosilases/genética , Neoplasias/genética , Sequência de Aminoácidos , Núcleo Celular/metabolismo , DNA Glicosilases/metabolismo , Reparo do DNA/fisiologia , Humanos , Isoenzimas/genética , Isoenzimas/metabolismo , Mitocôndrias/metabolismo , Dados de Sequência Molecular , Neoplasias/etiologia , Neoplasias/metabolismoRESUMO
Some beetles in the Namib Desert collect drinking water from fog-laden wind on their backs. We show here that these large droplets form by virtue of the insect's bumpy surface, which consists of alternating hydrophobic, wax-coated and hydrophilic, non-waxy regions. The design of this fog-collecting structure can be reproduced cheaply on a commercial scale and may find application in water-trapping tent and building coverings, for example, or in water condensers and engines.
Assuntos
Besouros/fisiologia , África Austral , Animais , Clima Desértico , ÁguaRESUMO
The most intense colours displayed in nature result from either multilayer reflectors or linear diffraction gratings. Here we investigate the spectacular iridescence of a spine (notoseta) from the sea mouse Aphrodita sp. (Polychaeta: Aphroditidae). The spine normally appears to be deep red in colour, but when light is incident perpendicular to the axis of the spine, different colours are seen as stripes running parallel to the axis of the spine; over a range of smaller incident angles, the complete visible spectrum is reflected with a reflectivity of 100% to the human eye. The simple structure responsible for this effect is a remarkable example of photonic engineering by a living organism.
Assuntos
Poliquetos/fisiologia , Animais , Medições Luminescentes , Óptica e Fotônica , Poliquetos/ultraestruturaRESUMO
This study reports a previously undescribed and unique sensillum. Some species of Lowrya Parker 1998 (Ostracoda: Cypridinidae) possess sensillae of an unusual type arranged in a single row parallel to, and near, the anterior, ventral and posterior margins of the external surface of each carapace valve. These sensilla are here termed coelotrichs, and are each centred within an almost spherical cavity in the carapace. The cavity forms at the base of a depression, and the depression in turn lies within a sunken channel on the exterior carapace surface. The opening of the coelotrich cavity bears a 'plug', through which only the widened, divided base of a seta passes. The seta is very fine and stiff throughout most of its length, and exhibits no pores. For most of its length the seta is orientated parallel to the carapace surface, shielded within the sunken channel. The coelotrich is probably a velocity detector, acted upon by the viscous drag of the surrounding fluid. Its function is probably to detect vibrations in the substrate, such as acoustic motion in the surrounding liquid, or steady fluid drainage motion.