Evidence for xylem feeding by Philaenus spumarius (L.) (Homoptera: Cercopidae)

Histological studies showed that the salivary sheaths of both nymphs and adults of P. spumarius penetrated principally to the xylem vessels of their host plants. Large amounts of liquid excrement were produced from one feeding site. The amount of excrement produced per unit body weight was much greater than for a phloem feeding aphid or psyllid.

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Zusammenfassung Philaenus spumarius ernährt sich von Xylemgefässsaft. Grosse Mengen von Ausscheidung wurden von einer einzigen Stichstelle aus gebildet, was auf Ernährung aus Gefässen, d.h. auf Extraktion von Phloem- oder Xylemsaft, hindeutet. Histologische Untersuchungen von Speichelscheiden zeigten, dass die meisten Einstichstellen in Xylemgefässen endigten. Der Inhalt der Parenchymzellen rings um die Speichelscheide war nicht entfernt und es kam nicht zu Nekrose. Larven und Adulte von P. spumarius nahmen 600 bis 1200 mal ihr eigenes Trockengewicht an Xylemsaft pro 24 Stunden auf. Dies ist viel mehr als bei Phloemsaugern. Wahrscheinlich handelt es sich um eine Konsequenz der geringeren Konzentration an Nährstoffen, wie Aminosäuren und Zucker, im Xylemsaft verglichen mit Phloemsaft.

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Supported by a Natural Environment Research Council studentship and supervised by Dr. J. B. Whittaker.     

Elytral polymorphism in spittlebugs: Philaenus spumarius (L.) from the Inner Hebrides

The frequencies of colour and pattern morphs of the elytra of Philaenus spumarius are similar on three of the Small Isles of the Inner Hebrides (Eigg, Canna and Muck); the population on the fourth island (Rhum: is somewhat distinct but is like that at sea level on the nearby mountainous mainland. These similarities and differences suggest that these morph frequencies are regulated by natural selection.     

Conditions requisite for stability of polymorphic balance in Philaenus spumarius (L.) (Homoptera)

Frequencies of colour morphs were recorded in 1969–74 in 5 island populations of Philaenus spumarius. The polymorphic balance was accurately maintained in three of them, but less constantly in the other two. One of the latter populations experienced a great fluctuation in size, and this was accompanied by a profound alteration in morph frequencies.One of the five populations consists of two subpopulations about 5 m apart. Although differing in the same island-specific respects from the populations of the other islands, the subpopulations were significantly unlike in the detailed distribution of the morph frequencies. These differences were probably due to the dissimilar distribution of the subniches available to the subpopulations. The dissimilar, island-specific patterns of morph frequences recorded in the other four populations may also be due to dissimilar subniche distribution. Migration between islands occurs at too slight a rate to cause an appreciable dysmetric load in the populations. In general, the rate of migration in Philaenus appears to be governed by the need for a balance between effective ability to colonize and effective subniche-specific protection of polymorphism.The study has been aided by grants from the University of Helsinki and from the National Research Council for Sciences, and the N.R.C. for Agriculture and Forestry (Academy of Finland).This paper is report no. 528 from Tvärminne Zoological station.     

Enzyme gene heterozygosity in small island populations of Philaenus spumarius (L.) (Homoptera)

Genetic polymorphism at about twenty enzyme loci in one mainland and in six differentially isolated island populations of Philaenus spumarius (L.) was studied by starch gel electrophoresis. The populations have different average degrees of heterozygosity, so that the most isolated population is the least polymorphic. The differences in heterozygosity seem to be correlated to the size of the population and the degree of isolation from other populations. With a single exception, the most common allele in each locus is the most common one everywhere. The results are compared with the differences observed in the color polymorphism of Philaenus island populations. The allele frequencies of enzyme loci are maintained by selection; the fact that the prevalent allele is the same in all populations may be due to selection and founder principle.Report no. 485 from the Tvärminne Zoological Station, University of Helsinki.     

The classification of adult colour forms of Philaenus spumarius (L.) (Homoptera: Insecta)

Philaenus spumarius is an insect of wide geographical distribution which has a number of colour morphs. This is a report of an attempt to provide an objective classification of the morphs by direct measurement of some visual characteristics. Distribution of these within the sample did not follow a normal curve but it was not possible to correlate the observed distribution with the subjectively sorted groups. This throws some doubt on the validity of the traditional subjective classification and on deductions based on this.     

Ventral pigmentation variation in the meadow spittlebug Philaenus spumarius (L.) (Hemiptera,Cicadomorpha: Cercopidae)

The meadow spittlebug Philaenus spumarius (L.) (Hemiptera, Cicadomorpha: Cercopidae) exhibits heritable colour/pattern variation on the dorsal and ventral sides of the adults. Eleven principal dorsal morphs are grouped as melanics and non-melanics according to the pigmentation of the dorsal surface. An investigation of ventral pigmentation variation in Philaenus spumarius was carried out with laboratory breeding stock obtained from Llysdinam, Wales. Analyses of the 1552 individuals, 753 females and 799 males, indicated that the pigmentation on the ventral surface of the adult individuals which vary significantly is associated with dorsal morph. Ventral parts of the dorsal melanics are usually darker than in non-melanics in both females (Kruskal-Wallis one-way ANOVA: H = 372.70, d.f. = 7, P < 0.001) and males (H = 407.36, d.f. = 7, P < 0.001). The combined “C” group morphs (flavicollis + gibbus + leuco- cephalus) are always darker than all other morphs in both sexes.     

Relationships between foliage and conducting xylem in Picea abies (L.) Karst.

Summary The relationship of leaf biomass and leaf area to the conductive area of stems and branches was investigated in Picea abies. A total of 30 trees were harvested to determine if these relationships were different in different crown zones and in trees growing with and without competition for light. Two methods were compared. In the first, data were accumulated from crown zones situated at the top of trees to the bottom; in the second, data were used from individual crown zones. The results indicated that the latter method is much more sensitive in detecting differences in the relationship of leaf biomass or leaf area to conductive area. The analysis also indicated that ratios such as leaf area/sapwood area are frequently size-dependent. This size-dependency can in some cases result in the differences being abscured, but more often leads to the false impression that the relationship between the variables changes. The relationship between leaf biomass and leaf area and conductive area of stems or branches was different in different crown zones and under different growth conditions. The slopes of these regressions appear to increase with decreasing transpirational demand and decrease with increasing hydraulic conductivity. The intercepts are probably related to the amount of identified sapwood actually involved in water conductance.     

Relationships between xylem sap constituents and leaf conductance of well-watered and water-stressed maize across three xylem sap sampling techniques

Many different techniques have been used for xylem sap collection, but few direct comparisons of techniques have been conducted and few comparisons have been based on comprehensive analyses of xylem sap. Moreover, the suitability of extraction techniques for use on plants grown under water-stress conditions has not been addressed. Xylem sap was extracted from both well-watered and water-stressed Zea mays plants using three different techniques. The main aim was to determine how the extraction method altered the correlations between sap constituents and stomatal conductance in order to determine which relationships change with extraction technique. A 'root pressure' technique was the simplest method of extracting large volumes of sap, but the low sap delivery rates altered the composition of sap. Two pressurization techniques that varied in the position from which sap was collected were tested. The pressurization techniques allowed for the control of delivery rates that influence sap constituent concentrations. The position from which xylem sap was collected on the plant was also found to be important. All three techniques produced consistent correlations between ABA and chloride delivery rates and changes in stomatal conductance, suggesting that each technique could be applied to identify certain putative xylem-borne signals.     

Post-embryonic photoreceptor development and dark/light adaptation in the spittle bug Philaenus spumarius (L.) (Homoptera, Cercopidae)

The aims of this paper have been to describe (1) the general structure of the compound eye of the spittle bug Philaenus spumarius, (2) the eye's post-embryonic development, (3) photomechanical changes upon dark/light adaptation in the eye, and (4) how leaving the semi-aquatic foam bubble and turning into an adult affects the organization of the eye. Spittle bugs, irrespective of size or sex, possess apposition type compound eyes. The eye's major components (i.e. facet, cornea, cone and rhabdom) grow rather isometrically from the smallest nymph to the adult. Photomechanical changes can occur during both nymphal and adult phases and manifest themselves through pigment granules and mitochondria migrating to and away from the rhabdom, and rhabdom diameters varying with time of day and ambient light level. When a nymph transforms into an adult, its compound eyes’ dorsoventral axes widen, facet diameters increase, facet shapes turn from circular to pentagonal and hexagonal, the cornea thickens and the rhabdoms become thinner. The agile adults, free from the foam that surrounds the nymphs, can be expected to need their vision more than the nymphs, and the changes in eye structure do, indeed, indicate that the adults have superior visual acuity. A thicker cornea in the adults reduces water loss and protects the compound eye from mechanical and light-induced damage: protection given to the nymphs by their foam bubbles.     

Identification of the silicon form in xylem sap of rice (Oryza sativa L.)

Rice (Oryza sativa L.) is a typical silicon (Si)-accumulating plant, but the mechanism responsible for the translocation from the root to the shoot is poorly understood. In this study, the form of Si in xylem sap was identified by (29)Si-nuclear magnetic resonance (NMR) spectroscopy. In rice (cv. Oochikara) cultured in a monosilicic acid solution containing 0.5 mM Si, the Si concentration in the xylem reached 6 mM within 30 min. In the (29)Si-NMR spectra of the xylem sap, only one signal was observed at a chemical shift of -72.6 ppm, which is consistent with that of monosilicic acid. A (1)H-NMR study of xylem sap did not show any significant difference between the wild-type rice and mutant rice defective in Si uptake, and the components of the xylem sap were not affected by the Si supply. The Si concentration in the xylem sap in vitro decreased from an initial 18 mM to 2.6 mM with time. Addition of xylem sap to a solution containing 8 mM Si did not prevent the polymerization of silicic acid. All these results indicate that Si is translocated in the form of monosilicic acid through the xylem and that the concentration of monosilicic acid is high in the xylem only transiently.     

The spittlebug Philaenus spumarius feeds from mature xylem at the full hydraulic tension of the transpiration stream

Feeding dynamics in the spittlebug Philaenus spumarius were analysed to determine the food source. Evidence from previous reports indicates that this species and similar insects feed from the xylem. However, it is not clear whether they extract sap from the main transpiration stream, or from some subsidiary compartment of xylem, such as immature or cavitated vessels. In feeding from such subsidiary compartments, the insects could avoid the large hydraulic tensions thought to prevail in xylem on the main transpiration stream. Philaenus , for example, is shown here to initiate feeding, and to feed rapidly, from plants in which the full xylem tension is estimated at −10 bar. From measurements of feeding rates and feeding patterns in stressed and unstressed plants, it is shown that Philaenus cannot feed from subsidiary xylem compartments. It is concluded that this insect feeds directly from the main transpiration stream, and at the full hydraulic tension which prevails there. The anatomy of the feeding pump in Philaenus appears consistent with this conclusion. Spittlebugs therefore offer an experimental tool for quasi- non-invasive sampling of the transpiration stream.     

Localized industrial melanism in the spittlebug Philaenus spumarius (L.) (Homoptera: Aphrophoridae) in Cardiff docks, south Wales

Twenty-four large adult samples of the meadow spittlebug, Philaenus spumarius , were collected from within a 4 times 3 km area containing Cardiff docks, south Wales. All samples are highly polymorphic for colour and pattern and show marked geographic variation in the combined frequency of the eight melanic morphs (from <15% to > 75%). Maximum melanic phenotype frequencies in both sexes coincide with dock wharfs currently containing large accumulations of pulverized coal and these samples are heavily contaminated with small coal particles. However, interpretation of this association is complicated by the existence, until 1976, of a smokeless fuel factory causing local air pollution immediately adjacent to the area of highest melanic frequency. It is suggested that the association of high levels of melanism in P. spumarius with areas heavily contaminated with particulate pollutants is due to direct selective effects of small particles favouring melanies.     

Effect of partial rootzone drying on the concentration of zeatin-type cytokinins in tomato (Solanum lycopersicum L.) xylem sap and leaves

Decreased cytokinin (CK) export from roots in drying soil might provide a root-to-shoot signal impacting on shoot physiology. Although several studies show that soil drying decreases the CK concentration of xylem sap collected from the roots, it is not known whether this alters xylem CK concentration ([CK(xyl)]) in the leaves and bulk leaf CK concentration. Tomato (Solanum lycopersicum L.) plants were grown with roots split between two soil columns. During experiments, water was applied to both columns (well-watered; WW) or one (partial rootzone drying; PRD) column. Irrigation of WW plants aimed to replace transpirational losses every day, while PRD plants received half this amount. Xylem sap was collected by pressurizing detached leaves using a Scholander pressure chamber, and zeatin-type CKs were immunoassayed using specific antibodies raised against zeatin riboside after separating their different forms (free zeatin, its riboside, and nucleotide) by thin-layer chromatography. PRD decreased the whole plant transpiration rate by 22% and leaf water potential by 0.08 MPa, and increased xylem abscisic acid (ABA) concentration 2.5-fold. Although PRD caused no detectable change in [CK(xyl)], it decreased the CK concentration of fully expanded leaves by 46%. That [CK(xyl)] was maintained and not increased while transpiration decreased suggests that loading of CK into the xylem was also decreased as the soil dried. That leaf CK concentration did not decline proportionally with CK delivery suggests that other mechanisms such as CK metabolism influence leaf CK status of PRD plants. The causes and consequences of decreased shoot CK status are discussed.     

Chemical forms of aluminum in xylem sap of tea plants (Camellia sinensis L.)

To identify the chemical forms of aluminum (Al) transported from roots to shoots of tea plants (C. sinensis L.), ²⁷Al-nuclear magnetic resonance and ¹⁹F NMR spectroscopy were used to analyze xylem sap.The concentration of Al in collected xylem sap was 0.29 mM, twice as high as that of F. Catechins were not detected in xylem sap. The concentration of malic acid in xylem sap was higher than that of citric acid, whereas the concentration of oxalic acid was negligible.There were two signals in the ²⁷Al NMR spectra of xylem sap, a larger signal at 11 ppm and a smaller one at −1.5 ppm. The former signal was consistent with the peak for an Al-citrate model solution, suggesting that an Al-citrate complex was present in xylem sap. Although the latter signal at −1.5 ppm was thought to indicate the presence of an Al-F complex (at 1.7 ppm) in xylem sap, there was only one signal at −122 ppm in the ¹⁹F NMR spectrum of xylem sap, indicating that the main F complex in xylem sap was F⁻.These results indicate that Al might be translocated as a complex with citrate, while Al-malate, Al-oxalate and Al-F complexes are not major Al complexes in xylem sap of tea plants.     

Novel approaches for examining the effects of differential soil compaction on xylem sap abscisic acid concentration, stomatal conductance and growth in barley (Hordeum vulgare L.)

Novel techniques were devised to explore the mechanisms mediating the adverse effects of compacted soil on plants. These included growing plants in: (i) profiles containing horizons differing in their degree of compaction and; (ii) split-pots in which the roots were divided between compartments containing moderately (1·4 g cm ^{? 3}) and severely compacted (1·7 g cm ^{? 3}) soil. Wild-type and ABA-deficient genotypes of barley were used to examine the role of abscisic acid (ABA) as a root-to-shoot signal. Shoot dry weight and leaf area were reduced and root : shoot ratio was increased relative to 1·4 g cm ^{? 3} control plants whenever plants of both genotypes encountered severely compacted horizons. In bartey cultivar Steptoe, stomatal conductance decreased within 4 d of the first roots encountering 1·7 g cm ^{? 3} soil and increased over a similar period when roots penetrated from 1·7 g cm ^{? 3} into 1·4 g cm ^{? 3} soil. Conductance was again reduced by a second 1·7 g cm ^{? 3} horizon. These responses were inversely correlated with xylem sap ABA concentration. No equivalent stomatal responses occurred in Az34 (ABA deficient genotype), in which the changes in xylem sap ABA were much smaller. When plants were grown in 1·7 : 1·4 g cm ^{? 3} split-pots, shoot growth was unaffected relative to 1·4 g cm ^{? 3} control plants in Steptoe, but was significantly reduced in Az34. Excision of the roots in compacted soil restored growth to the 1·4 g cm ^{? 3} control level in Az34. Stomatal conductance was reduced in the split-pot treatment of Steptoe, but returned to the 1·4 g cm ^{? 3} control level when the roots in compacted soil were excised. Xylem sap ABA concentration was initially higher than in 1·4 g cm ^{? 3} control plants but subsequently returned to the control level; no recovery occurred if the roots in compacted soil were left intact. Xylem sap ABA concentration in the split-pot treatment of Az34 was initially similar to plants grown in uniform 1·7 g cm ^{? 3} soil, but returned to the 1·4 g cm ^{? 3} control level when the roots in the compacted compartment were excised. These results clearly demonstrate the involvement of a root-sourced signal in mediating responses to compacted soil; the role of ABA in providing this signal and future applications of the compaction procedures reported here are discussed.     

The chemical form of trivalent chromium in xylem sap of maize (Zea mays L.)

Abstract

The toxicity, mobility and bioavailability of Cr, a versatile industrial metal and a contaminant, depends on its chemical form, viz: Cr(lll) and Cr(VI). It may enter humans through plants grown on contaminated soil or irrigated by contaminated water. The phytoavailability and transfer through agricultural food chains requires an understanding of mechanisms of Cr uptake and translocation by plants. Xylem sap transports both nutrient and non-nutrient ions after absorption by roots to aerial parts of the plant. lt transports cations by complexation with organic ligands. Trivalent chromium, though prone to hydrolysis, also complexes O donor ligands. The chemical form in which Cr(lll) is transported by xylem sap was investigated. ln vitro studies were performed by mixing the xylem sap of maize plants at three stages of plant growth with radiotagged Cr(III). The speciation change was investigated after 10 days and 30 days by anion and cation exchange elution chromatography. The elution curves were compared with those of pure Cr(III) and Cr(III) complexes of different synthetic acids. Complexation of Cr(III) with ligands of xylem sap especially with carboxylates was evident. Cationic Cr(III) was vitally being transported as anionic organic complex species. The major species seemed to be that of Cr(III)-citrate. Citric acid was the major complexing acid of xylem sap as determined by HPLC. These mobile and soluble complexes may get immobilized and stored in leaves and other edible plant parts. This may also be a mechanism used by plants for detoxification of toxic Cr(VI) which may become reduced and then complexed.     

The presence of N-acetylneuraminic acid in Malpighian tubules of larvae of the cicada Philaenus spumarius

Sialic acid-containing glycoconjugates are generally considered to be unique to the deuterostomes, a lineage of the animal kingdom which includes animals from the echinoderms up to the vertebrates. There are, however, two isolated reports of sialic acid occurring in the insect species Drosophila melanogaster and Galleria mellonella. Since insects are classified as protostomes, these findings call previous assumption on the phylogenetic distribution and thus on the evolution of sialic acids into question. Here, we report the occurrence of N-acetylneuraminic acid (Neu5Ac) in larvae of the cicada Philaenus spumarius. Cytochemical analysis of larval sections with lectins from Sambucus nigra and Limax flavus suggested the presence of sialic acids in the concrement vacuoles of the Malpighian tubules. The monoclonal antibody MAb 735, which is specific for polysialic acid, labelled the same structures. A chemical analysis performed by HPLC of fluorescent derivatives of sialic acids and by GLC-MS provided sound evidence for the presence of Neu5Ac in the Philaenus spumarius larvae. These data suggest that in this cicada Neu5Ac occurs in 2,8-linked polysialic acid structures and in 2,6-linkages. The results provide further evidence for the existence of sialic acids in insects and in linkages known to occur in glycoconjugates of deuterostomate origin.     

Temperature and the colour polymorphism of Philaenus spumarius (Homoptera: Aphrophoridae)

ABSTRACT.

• 1 The spittle bug, Philaenus spumarius (L.), has eleven distinct elytral colour pattern morphs in Britain, of which three are designated non-melanic and eight melanic. Frequencies of different morphs vary between populations.
• 2 The role of thermal factors in the maintenance of the balanced polymorphism was investigated by measurements of dorsal reflectance and body temperature when artifically irradiated.
• 3 Non-melanic forms had higher reflectances and showed lower temperature excesses when irradiated. There was considerable variation in reflectance for the populi–typicus morph group, and trilineatus is therefore the only true‘non-melanic’morph.
• 4 For samples along altitudinal transects in three Scottish glens, trilineatus frequencies were found to be negatively and populi-typicus and melanic frequencies positively correlated with altitude.
• 5 Thermal considerations do therefore play some role in the broad and complex selection regime of P.spumarius.

     

Inhibitor content of phloem and xylem sap obtained from willow (Salix viminalis L.) entering dormancy

Summary It is shown that there is one growth inhibitor in the phloem sap and two in the xylem sap of willow (S. viminalis L.). The concentration of the inhibitor in the phloem sap, (+)-abscisic acid, increases as the plants enter dormancy. This is also shown for (+)-abscisic acid in the xylem sap, but the concentration of the second inhibitor decreases in a reciprocal manner.