“Lethal effects of the wild potato Solanum bulbocastanum on the planthopper Pentastiridius leporinus, a vector of bacterial pathogens in potato”

“Lethal effects of the wild potato Solanum bulbocastanum on the planthopper Pentastiridius leporinus, a vector of bacterial pathogens in potato” | bioRxiv /* */ /* */ <!-- <!-- /*! * yepnope1.5.4 * (c) WTFPL, GPLv2 */ (function(a,b,c){function d(a){return"[object Function]"==o.call(a)}function e(a){return"string"==typeof a}function f(){}function g(a){return!a||"loaded"==a||"complete"==a||"uninitialized"==a}function h(){var a=p.shift();q=1,a?a.t?m(function(){("c"==a.t?B.injectCss:B.injectJs)(a.s,0,a.a,a.x,a.e,1)},0):(a(),h()):q=0}function i(a,c,d,e,f,i,j){function k(b){if(!o&&g(l.readyState)&&(u.r=o=1,!q&&h(),l.onload=l.onreadystatechange=null,b)){"img"!=a&&m(function(){t.removeChild(l)},50);for(var d in y[c])y[c].hasOwnProperty(d)&&y[c][d].onload()}}var j=j||B.errorTimeout,l=b.createElement(a),o=0,r=0,u={t:d,s:c,e:f,a:i,x:j};1===y[c]&&(r=1,y[c]=[]),"object"==a?l.data=c:(l.src=c,l.type=a),l.width=l.height="0",l.onerror=l.onload=l.onreadystatechange=function(){k.call(this,r)},p.splice(e,0,u),"img"!=a&&(r||2===y[c]?(t.insertBefore(l,s?null:n),m(k,j)):y[c].push(l))}function j(a,b,c,d,f){return q=0,b=b||"j",e(a)?i("c"==b?v:u,a,b,this.i++,c,d,f):(p.splice(this.i++,0,a),1==p.length&&h()),this}function k(){var a=B;return a.loader={load:j,i:0},a}var l=b.documentElement,m=a.setTimeout,n=b.getElementsByTagName("script")[0],o={}.toString,p=[],q=0,r="MozAppearance"in l.style,s=r&&!!b.createRange().compareNode,t=s?l:n.parentNode,l=a.opera&&"[object Opera]"==o.call(a.opera),l=!!b.attachEvent&&!l,u=r?"object":l?"script":"img",v=l?"script":u,w=Array.isArray||function(a){return"[object Array]"==o.call(a)},x=[],y={},z={timeout:function(a,b){return b.length&&(a.timeout=b[0]),a}},A,B;B=function(a){function b(a){var a=a.split("!"),b=x.length,c=a.pop(),d=a.length,c={url:c,origUrl:c,prefixes:a},e,f,g;for(f=0;f<d;f++)g=a[f].split("="),(e=z[g.shift()])&&(c=e(c,g));for(f=0;f<b;f++)c=x[f](c);return c}function g(a,e,f,g,h){var i=b(a),j=i.autoCallback;i.url.split(".").pop().split("?").shift(),i.bypass||(e&&(e=d(e)?e:e[a]||e[g]||e[a.split("/").pop().split("?")[0]]),i.instead?i.instead(a,e,f,g,h):(y[i.url]?i.noexec=!0:y[i.url]=1,f.load(i.url,i.forceCSS||!i.forceJS&&"css"==i.url.split(".").pop().split("?").shift()?"c":c,i.noexec,i.attrs,i.timeout),(d(e)||d(j))&&f.load(function(){k(),e&&e(i.origUrl,h,g),j&&j(i.origUrl,h,g),y[i.url]=2})))}function h(a,b){function c(a,c){if(a){if(e(a))c||(j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}),g(a,j,b,0,h);else if(Object(a)===a)for(n in m=function(){var b=0,c;for(c in a)a.hasOwnProperty(c)&&b++;return b}(),a)a.hasOwnProperty(n)&&(!c&&!--m&&(d(j)?j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}:j[n]=function(a){return function(){var b=[].slice.call(arguments);a&&a.apply(this,b),l()}}(k[n])),g(a[n],j,b,n,h))}else!c&&l()}var h=!!a.test,i=a.load||a.both,j=a.callback||f,k=j,l=a.complete||f,m,n;c(h?a.yep:a.nope,!!i),i&&c(i)}var i,j,l=this.yepnope.loader;if(e(a))g(a,0,l,0);else if(w(a))for(i=0;i (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0];var j=d.createElement(s);var dl=l!='dataLayer'?'&l='+l:'';j.src='//www.googletagmanager.com/gtm.js?id='+i+dl;j.type='text/javascript';j.async=true;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-M677548'); Skip to main content Home About Submit ALERTS / RSS Search for this keyword Advanced Search New Results “Lethal effects of the wild potato Solanum bulbocastanum on the planthopper Pentastiridius leporinus , a vector of bacterial pathogens in potato” View ORCID Profile Eva Therhaag , View ORCID Profile Karin Gorzolka , View ORCID Profile Jürgen Gross doi: https://doi.org/10.1101/2025.10.12.681902 Eva Therhaag 1 Julius Kühn Institute (JKI) - Federal Research Centre for Cultivated Plants, Institute for Plant Protection in Fruit Crops and Viticulture , Schwabenheimer Str. 101, 69221 Dossenheim, Germany 2 Geisenheim University, Department of Crop Protection , Von-Lade-Str. 1, 65366 Geisenheim, Germany Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Eva Therhaag Karin Gorzolka 3 Julius Kühn Institute (JKI) - Federal Research Centre for Cultivated Plants, Institute for Ecological Chemistry, Plant Analysis and Stored Product Protection , Königin-Luise-Straße 19, 14195 Berlin, Germany Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Karin Gorzolka Jürgen Gross 1 Julius Kühn Institute (JKI) - Federal Research Centre for Cultivated Plants, Institute for Plant Protection in Fruit Crops and Viticulture , Schwabenheimer Str. 101, 69221 Dossenheim, Germany 2 Geisenheim University, Department of Crop Protection , Von-Lade-Str. 1, 65366 Geisenheim, Germany Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Jürgen Gross For correspondence: juergen.gross{at}julius-kuehn.de Abstract Full Text Info/History Metrics Supplementary material Preview PDF Abstract The planthopper Pentastiridius leporinus [Hemiptera: Cixiidae] is a vector of the two plant pathogenic bacteria ‘ Candidatus Arsenophonus phytopathogenicus’ and ‘ Candidatus Phytoplasma solani’ in potato. An infection leads to symptoms such as rubbery and non-processable tubers and consequently to substantial decreases in yield and tuber quality as well as to abandonment of seed potato production in affected areas. Up to date, resistant potato varieties are not yet known, and the tool box of integrated pest management to prevent the spread of P. leporinus as causal vector is quite limited. Mortality experiments designed as no-choice trials on potato cultivar ‘Gala’ ( Solanum tuberosum ) and on its wild relative S. bulbocastanum revealed significant effects of the latter on the survival rate of the vector. LC/MS analysis of the insects and their intestines showed that acyl solamines were retrieved only in individuals from the S. bulbocastanum plants. Furthermore, choice-tests were carried out to study the vector behavior. A preference towards S. bulbocastanum vs. S. tuberosum ‘Gala’ was observed despite its lethal effects. Volatile organic compounds of the two different species were collected and analyzed by GC/MS. The two profiles differed in 39 of 80 compounds. The wild potato species S. bulbocastanum and its compounds are discussed as potential starting points for further research on sustainable management of P. leporinus . Key Message We tested a wild potato against a planthopper which vectors pathogenic bacteria to potato Wild potato led to higher mortality of the planthopper but was preferred against cultivated potato Uptake assays suggest that the phloem sucking planthopper is taking up xylem sap as well Acyl solamines were only found in guts from planthoppers sucking on the wild potato Our results present a starting point for new approaches for sustainable vector control 1 Introduction The history of potato ( Solanum tuberosum L.) cultivation dates back 7,000 years. Potato belongs to the first domesticated food crop and still plays an important role in human diets and nutrition – it is grown on each continent despite Antarctica ( Zaheer and Akhtar 2016 ; Bakhsh et al. 2023 ). In 2023, the global potato production was estimated about 383 million tons ( FAO 2024 ). A bit more than a twelfth of that, 48.3 million tons of potatoes were harvested in Europe. There, the five countries Germany, France, the Netherlands, Poland and Belgium made up almost 70% of the harvested area ( Eurostat 2024 ). Since 2022, the potato production in the Southwest of Germany encounters severe challenges. In particular, in Hesse, the production of seed potato decreased dramatically over the last few years from 290 ha in 2021 to 67 ha in 2024 ( LLH 2024 ). This coincidences with the first finding of a relatively new vector in this crop, the planthopper Pentastiridius leporinus L. [Hemiptera: Cixiidae] ( Behrmann et al. 2023 ). It transmits the bacterial pathogens ‘ Candidatus Phytoplasma solani’ and ‘ Candidatus Arsenophonus phytopathogenicus’ to potato ( Behrmann et al. 2023 ; Therhaag et al. 2024a ). These transmitted pathogens have been reported recently as an emerging risk in a stakeholder discussion meeting of the European Food Safety Agency ( EFSA 2024 ). An infestation of the crop may be associated with symptoms of wilting, yellow to purplish leaf tops, rubbery, small or airy tubers and loss of yield ( Therhaag et al. 2024a ; Rinklef et al. 2024 ). In tubers, infection with the bacteria leads to crop losses, partly due to reduced storage, processing or germination capacity ( Lindner et al. 2011 ; Mahillon et al. 2025 ; Lang et al. 2025 ). Since adult P. leporinus have only been reported quite recently as vector of the afore mentioned pathogens in potato ( Therhaag et al. 2024a ), specific vector management options in potatoes have not yet been studied. In 2024, other crops and vegetables have been reported to be infested by one or both of the transmitted pathogens ( Therhaag et al. 2024b ; Lang et al. 2025 ), which aggravates the situation in areas of high abundance of P. leporinus . This underlines the urgent need to investigate and develop short- and long-term management tools. Since the planthopper is well adapted to the cropping rotation, which includes winter cereals sown after potato or sugar beet, a starvation period due to fallow, maize or other summer crops may be a promising attempt to break its lifecycle and reduce the population ( Bressan et al. 2010 ; Pfitzer et al. 2024 ). An intercrop that has a negative impact on the planthopper could also be included in the crop rotation. Besides the lever of an adapted crop rotation, another important part of integrated pest management (IPM) is the use of resistant or robust varieties, in particular when bacterial plant pathogens are transmitted by polyphagous vectors across different crops ( Agrios 2005 ). In the case of potato, various wild relatives are available which could serve as a source of desirable traits such as resistance or resilience to better cope with pests and diseases ( Jansky and Spooner 2018 ). Efforts have been made to introduce resistance genes from Solanum species to domesticated potato as for example reported for potato late blight caused by Phytophthora infestans ( Malcolmson and Black 1966 ; Van Der Vossen et al. 2003 ; Gopal 2023 ) or the Colorado potato beetle ( Leptinotarsa decemlineata ) ( Crossley et al. 2018 ). However, no resistance traits against P. leporinus or the transmitted pathogenic bacteria were found in cultivated or wild potato species until today. Recently, bioassays with a bioactive compound isolated from the wild relative Solanum bulbocastanum Dun. showed effects against the Colorado potato beetle ( L. decemlineata ), the potato flea beetle ( Epitrix papa ), and P. infestans ( Gorzolka et al. 2025 ). The isolated compound (C16:3(7Z,10Z,13Z)-solamine) is an acyl polyamine. An acyl solamine fraction obtained from S. bulbocastanum leaf material with about 90% of C16:3(7Z,10Z,13Z)-solamine inhibited the growth of the necrotrophic pathogens Alternaria solani and Botrytis cinerea ( Lenz et al. 2025 , accepted). In the present study, we investigated for the first time potential effects of S. bulbocastanum on the planthopper P. leporinus in comparison with the commercial potato cultivar ‘Gala’. In no-choice and choice bioassays we investigated both the preferences of the planthopper and its mortality after feeding on the two test plants. We analyzed the content of planthoppers and their gastro-intestines by LC-MS for proof of the cause of their death. Furthermore, the volatile compounds of the wild relative and the potato cultivar were analyzed by GC-MS to find out, if the odors may differ or could act as olfactory cues to the planthopper. We discuss S. bulbocastanum as a potential trap plant for P. leporinus , the role of its bioactive compounds and its use in resistance breeding as medium to long-term management options to control P. leporinus along with other important potato pests and diseases. 2 Material and Methods 2.1 No-choice mortality experiments Planthoppers Two no-choice mortality experiments took place subsequently at the Julius Kühn Institute in Dossenheim, Germany. For the first no-choice experiment, planthoppers were collected on 24 June 2024 in Southwestern Germany with sweeping nets at different sites (potato fields in Bingen 49°56’52.8’’N 7°56’03.6’’E, and Frankenthal 49°33’03.9’’ 8°23’22.1’’E, winter wheat field in Bürstadt 49°38’17.5’’N 8°24’57.6’’E). Likewise, planthoppers were collected in Bad Wimpfen (sugar beet field 49° 13’ 18.7’’N 9° 8’ 36’’E) on 15 July 2024 for the repeated no-choice mortality experiment. The planthoppers have been stored in insect cages and transported to the experiment site. There, ten to eleven individuals have been placed in each of eight transmission cylinders planted with S. tuberosum cultivar ‘Gala’, and S. bulbocastanum . Specimen of P. leporinus only were used after morphological identification (Biedermann, Robert and Niedringhaus, Rolf 2009). The experiments started the same day of the field collection. Both experiments took place in the same climate chamber at 22 °C, 55% relative humidity, and a light/dark cycle of 18/6 h. Insects that died during the experiment were removed daily and stored at −20 °C until further analysis. Same applied to the remaining insects at the end of the trial, i.e. after 11 days. Plant material S. tuberosum cultivar ‘Gala’ (Julius Kühn Institute (JKI), Institute for Breeding Research on Agricultural Crops, Sanitz, Germany) was planted in 3L pots filled with potting soil and sand in a ratio of 40:10 L on 3 June 2024. One tuber per pot was planted. Equal planting applied to S. bulbocastanum WKS31741, clonal line blb2G (WKS31741 seed material obtained from Federal ex situ Gene Bank of the Leibniz Institute of Plant Genetics and Crop Plant Research, Gatersleben, Germany), clonal line maintained from JKI (Sanitz, Germany), which resulted in one shoot per pot. All plants were grown in an insect-free greenhouse. The same S. bulbocastanum plants were used for the first and the second experiment since this species grew slowly and tuber material was rare. In the case of ‘Gala’, all shoots but one were cut at emerging stage to fit inside the transmission cylinder ( Therhaag et al. 2024a ) for the first experiment. For the second experiment, a new shoot from the same plant which emerged on 5 July 2025 was taken. 2.2 Dual-Choice experiments Two subsequent experiments were conducted with both plant species in the same climate chamber with equal parameters as described above, starting on 5 and 10 July 2024, respectively, and merged to a choice experiment with a total of 12 replications. One pot with S. bulbocastanum and one pot with S. tuberosum were put into each of the insect cages (BugDorm 2S120, MegaView Science Co., Ltd., Taiwan). Planthoppers were collected in Southwestern Germany at the same site as described earlier (Bad Wimpfen) on 5 July 2024 and on 9 July 2024. Eight female and two male specimens of P. leporinus were placed in the middle of the cages at the same distance to both plants. The number of planthoppers on each plant was counted after 30 minutes, one hour, two hours, three hours, one day and two days later (i.e. after 24 h and 48 h respectively). 2.3 Assay for staining plant and insect tissues Adults of P. leporinus from insect rearing on sugar beet variety ‘Danicia’, at the Julius Kühn Institute in Dossenheim, Germany, were taken for the assay of plant sap uptake of S. bulbocastanum and S. tuberosum cv ‘Gala’. Cut twigs of potato cultivar ‘Gala’ or S. bulbocastanum were inserted through a 1 cm thin foam plate inside Falcon tubes filled with 4 ml solution of 0.5% Brilliant Blue (CAS-No. 3844-45-9, Carl Roth GmbH, Germany). The twig was cut at the bottom and top before insertion to facilitate quick water transport by transpiration. Single individuals of P. leporinus , female or male, were transferred to the air-filled part of Falcon tubes. The plate between the solution and the potato leaves prevented the planthopper from falling into the solution. The tube was covered with a mesh for air circulation and stored at room temperature for two days. After two days, both the plant and the insects were removed from the tube. For proof of the uptake of blue-dyed water into the xylem of the plants, stems of random samples of each species were cut with razor blades, viewed under a stereo microscope (ZEISS Axioplan) and photographed with a mounted digital camera (Olympus ColorView IIIu FW) using the imaging software analySIS docu 5.0 (Olympus Soft Imaging Solutions GmbH). The insects were fixed with insect pins on dental wax in a petri dish and dissected in phosphate-buffered solution as described by Takeshita and Kikuchi (2020) . The gastro-intestine and its content were easily recognizable by the blue colorant and cut off the insect body ventrally. Each sample was photographed under a stereo microscope (SteREO Discovery.V12, Carl Zeiss Microscopy GmbH, Jena, Germany) equipped with a camera (Axiocam 305, Carl Zeiss Microscopy GmbH, Jena, Germany), then transferred to 10 µl methanol (LC-MS grade) in 1.5 ml reaction tubes and stored at −20 °C until further extraction. 2.4 LC/MS metabolite profiling Sample collection and extraction The LC/MS metabolite profiling of whole insects was obtained from planthoppers of the second no-choice mortality trial. For this, dead planthoppers were collected daily in 1.5 ml reaction tubes and stored at −20 °C. For the LC/MS metabolite profiling of gut extracts, individuals of the assay for plant sap uptake were dissected as described earlier (section 2.3). Crude extracts of whole insects were prepared as follows: A 5-mm steal bead and 50 µl 80% (v/v) methanol/water with 0.5% (v/v) formic acid were added to each insect. The sample was vigorously shaken for 5 min at 30 Hz and room temperature using a mixer mill (MM300, Retsch GmbH, Haan, Germany). After short centrifugation for sample precipitation, samples were treated with ultrasonic at room temperature for 5 min and then vigorously shaken for 5 min at 30 Hz again. After 15 minutes of centrifugation (13000 g, 22 °C), the supernatant was collected for LC-MS analysis. Crude extracts of insect intestinal samples were extracted by adding 20 µl 80% methanol/water with 0.5% formic acid. Samples tissues were not disrupted, only the intestinal content was targeted for analysis. For this, samples were shaken on a vertical rotary shaker for 10 min (1000 rpm, RT) and centrifuged for 10 min (13000 rpm, 22 °C). The supernatant was subjected to LC-MS analysis. LC/MS Extracts were analyzed on an Infinity 1290 series UHPLC system hyphenated to a 6550 iFunnel Q-TOF mass spectrometer (Agilent Technologies, Santa Clara, CA) via a dual jet stream electrospray ion source at the Julius Kühn Institute in Berlin, Germany as described ( Tais et al. 2021 ). Samples (injection volume 5 µL for whole insect analyses; 10 µl for insect intestinal analyses) were separated on an ACQUITY UPLC HSS T3 column (2.1 mm × 100 mm, particle size 1.8 µm, pore size 100 Å, Waters Corporation, Milford, MA) using 0.1% (v/v) formic acid in water and 0.1% formic acid in acetonitrile as eluent A and B, respectively. The binary gradient program at a flow rate of 500 µL/min was 0-1 min: isocratic 5% B, 1-15 min: linear to 60% B, 15-18 min: linear to 95% B, 18-19 min: isocratic 95% B, 19-21 min: isocratic 5% B. The column temperature was 40 °C; the autosampler temperature 6 °C. Eluting compounds were detected in positive mode from m/z 50-1700 (centroid mode, 3 spectra/s) in all samples and in negative ion mode in intact insect total extracts; instrument settings were applied as described by Tais et al. (2021) . Analysis of LC/MS data Untargeted metabolite profiling was performed using the R package “xcms” Release (3.20) with peak picking using the ‘centWave’ method with a signal to noise ratio of SN = 4, otherwise using default settings. Peaks were aligned without the need of retention time correction. ‘fillPeaks’ was neglected. All peaks, including potential adducts and isotopes from retention times between 0.6 to 14 min were included in statistics. Data were imported in MetaboAnalyst ( Xia et al. 2009 ; Pang et al. 2021 , 2022 ) for visualization and statistical analysis. Raw data inspection, compound spectrum extraction, compound database search and generation of molecular formulas were done using MassHunter Qualitative Analysis software (version B.07.00, Agilent Technologies); in-house libraries ( Gorzolka et al. 2025 ) and the technology platform “Metlin” ( Guijas et al. 2018 ) were used for compound annotation by exact m/z match (5 ppm) as well as retention time match for positive library hits. 2.5 GC/MS analysis of Volatile Organic Compounds (VOCs) Collection of VOCs of the potato cultivar ‘Gala’ and S. bulbocastanum VOCs of six plants of the potato cultivar ‘Gala’ and of six plants of S. bulbocastanum were collected on 30 July 2024 from plants of the second no-choice mortality experiment. For the collection, a single shoot of each sample plant was placed inside a plastic oven bag which was heated beforehand at 60 °C for 4 hours and cooled again. Each bag was connected to a headspace collection device (FLUSYS GmbH, Offenbach, Germany), which was developed by J. Gross ( Rid et al. 2016 ; Gross et al. 2019 ) and improved in the PurPest Project ( European Commission 2022 ) described by Karimi and Gross (2024) . The collection was implemented in an insect-proof greenhouse chamber. The VOCs were trapped in stainless steel tubes filled with Tenax TA35/60 sorbent (Markes, Neu-Isenburg, Germany) until further analysis. The air inside of empty plastic bags was collected and analyzed as a control to avoid identification of compounds not originating from the plants. Analysis of GC/MS data A subsequent chemical analysis of the VOCs was undertaken at the day of collection through automated thermodesorption (TurboMatrix™ ATD 650, PerkinElmer, Rodgau, Germany) and gas chromatography coupled with mass spectrometry at the Julius Kühn Institute in Dossenheim, Germany as described by Gallinger et al. (2020) . In our study, the VOC flow was split so that 30% of the flow were subjected to a full-scan mass spectrometer monitoring at a range of 50-350 m/z. Obtained data were analyzed with AMDIS software (Automated Mass spectral Deconvolution and Identification System, V. 2.71; National Institute of Standards and Technology NIST, Boulder, CO) according to a protocol by Gross et al. (2019) . In AMDIS, ion fragmentation patterns, retention times and retention indices (RI) were compared for the identification of the detected compounds with those of standard compounds that were introduced in the same system and stored in an in-house library. For compounds not included in the in-house library, matches of the NIST library (National Institute of Standards and Technology) and the NIST Chemistry WebBook (2025) were taken for further comparison. In that case, only RI from gas chromatography systems with similar technical parameters in terms of column diameter and length, stationary phase material and comparable temperatures were considered. Compounds identified with the use of authentic standards are treated as “identified”, those obtained without the use of authentic standards as “annotated”. Compounds that could not be identified are annotated as “unknown”. The identification criteria of the AMDIS software were as follows: the minimum match factor was set to 80% using retention index (RI) data, the RI window was set to 20 with the match factor penalties set at very low level with 20 for maximum penalty and 20 for “no RI in library”. Compounds occurring in less than two (out of twelve) samples were excluded from the analysis as well as the internal standard and known synthetic compounds. Relative proportions of each VOC were calculated by division of the peak area through the sum of the peak areas of all compounds per sample. 2.6 Statistical analysis Statistical analysis of the mortality experiments, the choice-experiments and the analysis of VOC profiles was performed in RStudio/2024.12.0+467 (“Kousa Dogwood” Release for windows, Posit Software, PBC) using R version 4.4.2 (R Core Team 2024. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ ). In case of the no-choice mortality experiments, a multivariate Cox regression analysis was performed using the coxph() function of the R ‘survival’ package ( Therneau et al. 2024 ) considering ‘treatment’, i.e. the two different plant species, and ‘pot’ as covariates to account for the grouping of individuals in the pots with the different plant species. Survival curves were created with the survfit() function of the same package. With regards to the choice experiments investigating plant preferences, the counted number of planthoppers per plant species in each cage was analyzed using a generalized linear mixed model approach with a zero-inflated Poisson distribution and log-link function of the ‘glmmTMB’ package ( Brooks et al. 2017 ; McGillycuddy et al. 2025 ). The number of individuals still alive at the time of counting was taken into account as an offset. The model was fitted with plant species as fixed effect, the cages and observation timings as random effects. Model fit parameters were obtained from the summary() function. An evaluation of the differences, i.e. the contrasts, between the marginal means of the contrasting variable ‘plant species’ at the different observation timings was conducted using the averaged contrast analysis of the ‘modelbased’ package ( Makowski et al. 2025 ) with the function estimate_contrasts(). The confidence interval level was set to 0.95, and the Holm-Bonferroni method for p-adjustments in frequentist multiple comparisons was applied. Results were visualized using the ‘ggplot2’ package in R ( Wickham 2016 ). For the VOC profiles of the both species, a non-metric multidimensional scaling using Bray-Curtis dissimilarities plot was obtained using metaNMDS from the ‘vegan’ package in R ( Brückner and Heethoff 2016 ; Oksanen et al. 2025 ). Of the same package, the envfit() function was applied to the compositional data set with permutations set to 10,000. Anderson’s permutational multivariate analysis of variance and of dispersions ( Anderson 2001 ; Anderson et al. 2006 ) has been computed in R with PERMANOVA and PERMDISP from the Bray-Curtis dissimilarity matrix of the VOCs as described in Brückner and Heethoff (2016) . 3 Results 3.1 No-choice mortality experiments Two no-choice experiments were conducted with field-caught adults of P. leporinus which were placed inside cylinders with either S. tuberosum cv ‘Gala’ or S. bulbocastanum plants. For a period of 11 days, dead planthoppers were counted and collected daily. Analysis of mortality was undertaken with the number of dead planthoppers per day and plant species. Mortality increased significantly for planthoppers feeding on S. bulbocastanum revealed by several distinct significance tests ( Table 1 ). Both experiments showed that the effect of the grouping in ‘pot’ fails significance whereas the effect of the ‘treatment’, namely the effect of the different plant species exposed to the planthoppers, contributes significantly to the hazard ratio of death of the planthoppers of the studies. Survival plots of the planthoppers of both no-choice experiments are compared in Figure 1 and visualize impact of the plant species on the survival. View this table: View inline View popup Download powerpoint Table 1. Results of statistical tests of a multivariate Cox regression analysis of both no-choice experiments. The covariate ‘treatment’ refers to the two different plant species, i.e. S. bulbocastanum and the potato cv ‘Gala’, whereas the covariate ‘pot’ accounts for the grouping of the planthoppers to the plant species potted in 16 and 12 pots in experiment I (collected from potato and winter wheat) and II (collected from sugar beet), respectively. Significance codes apply as follows: ‘***’ for p < 0.001, ‘**’ for p < 0.01, ‘*’ for p |z|) column represents the p-value associated with the value in the z value column. Download figure Open in new tab Fig. 1. Survival plots including the confidence interval (0.95) of (A) the first (adults collected from potato and winter wheat), and (B) the second (adults collected from sugar beet) no-choice experiment (B) with P. leporinus 3.2 Dual-Choice experiment In the dual-choice experiment, field-caught planthoppers were placed inside the insect cages and their total number was counted on each plant species at different observation times over a period of two days, with individuals still alive chosen as offset. As can be drawn from Figure 2 , planthoppers which were decisive for a plant, preferred the crop wild relative at the different observation timings. A marginal contrast analysis with a Holm-Bonferroni method for p-adjustments revealed significance levels ranging between p < 0.001 and p < 0.05 ( Table 2 ) in the chosen confidence interval of 95%. View this table: View inline View popup Download powerpoint Table 2. Results of the average contrast analysis of the choice experiment displaying the differences of the marginal means of the contrasting variable ‘plant species’ at different observation timings, with p1: S. bulbocastanum , p2: S. tuberosum cv ‘Gala, SE: standard error, z: z-value, df: degrees of freedom, and t1 to t6: 0.5h, 1h, 2h, 3h, 24h, 48h. P-values are obtained from the Holm-Bonferroni method with significance codes as follows: ‘*’ for p < 0.05, ‘**’ for p < 0.01, and ‘***’ for p < 0.001 within the 95% confidence interval (95% CI) Download figure Open in new tab Fig. 2. Preference of planthoppers for a plant species S. bulbocastanum or S. tuberosum cv ‘Gala’ observed in cages ( n =12) of the choice experiment. Boxplots represent the percentage of observed planthoppers related to the number of planthoppers still alive; bars indicate standard error (SE); at the observation timing 1d, only data of six cages ( n =6) are available; asterisks indicate significant adjusted p-values obtained from the Holm-Bonferroni method with ‘*’ for p < 0.05, ‘**’ for p < 0.01, and ‘***’ for p < 0.001, in the 95% confidence interval 3.3 Assay for plant sap uptake For the LC/MS analysis of planthopper intestines, blue dyed water was used for plant and eventually insect uptake to facilitate recognition and excision of the intestines. Photographs of petiole sections show some cells with blue content, presumably xylem cells stained by uptake of blue colored water by transpiration ( Figure 3 A ). Photographs of dissected insects show the blue stained intestine could be easily distinguished from other organs ( Figure 3 B ). Download figure Open in new tab Fig. 3. A: Photograph of a section of a cut potato leaf petiole after insertion of the twig in blue-dyed water for two days (25x magnification); red arrow points to cells of the vascular bundle filled blue. B: Stacked photograph before excision of intestine with blue content of female P. leporinus , 15.2x magnified 3.4 LC/MS analysis The untargeted LC/MS-analysis of a) the extracts of the whole insect bodies, and b) the extracts of intestines of the planthoppers resulted in 2,544 and 1,976 signals with retention times between 0.6 min to 14 min, respectively. A fivefold-change analysis was calculated to compare the absolute value of change between the two group means of planthoppers which fed on S. bulbocastanum vs. S. tuberosum cv ‘Gala’. An adjusted p-value (FDR) threshold of p<0.05 was taken for the unpaired t-tests. Figure 4 shows a volcano plot combining the fold change analysis and the t-test values of the detected compounds with the acyl solamine signals labelled. This analysis reveals that out of the 2,544 analyzed signals in the insect bodies, only C16:3-solamine [M+H]+ was significantly more than 5-fold enhanced in insects that fed on S. bulbocastanum with p<0.05. Of the 1,976 detected compounds of the intestine extracts, also acyl solamine signals (C16:3-solamine [M+H]+, C16:3-solamine [M+H]2+) had the highest fold change enhancement and significance level in S. bulbocastanum derived planthoppers. The significantly changing metabolites of both datasets are listed in Table 3 . View this table: View inline View popup Download powerpoint Table 3. Fold change and significance analyses of metabolite profiling data obtained from LC-MS extracts of A) whole plant hopper with in total 2,544 signals and B) plant hopper intestines with in total 1,976 signals. Thresholds were 5-fold change (FC) and FDR-adjusted p-values of p<0.05. Identified compounds are indicated in “annotation”. Signals “Unknown 3 Isotope 1-3” derived from one single metabolite. C16:3-sol: C16:3-solamine. A fold change of 1 indicates enhanced abundances in Solanum tuberosum derived plant hopper samples. “Unknown”: no match in data bases Download figure Open in new tab Fig. 4. Volcano plots of untargeted metabolite profiling data with unpaired samples of extracts of whole insect bodies of P. leporinus (A) and extracts of intestines from planthoppers (B) fed on S. bulbocastanum vs. S. tuberosum cv ‘Gala’. The x-axis shows the fold change (FC) of the compound in log2 scale, the y-axis shows the significance of the fold change as −log10 (p-value) based on FDR adjusted p-values from the t-tests. Points indicate compounds; compounds enhanced in S bulbocastanum group on the left side, compounds enhanced in S. tuberosum cv ‘Gala’ on the right side. Vertical solid lines indicate thresholds of 5x fold change. Horizontal lines indicate a significance threshold of p=0.05, points above this line are significantly (p5 fold change with p<0.05 A Mann-Whitney U test was applied to the data of the C16:3-solamine peak area of the LC/MS analyses of extracts obtained from planthoppers either feeding on S. bulbocastanum or S. tuberosum cv ‘Gala’. With regards to extracts of whole planthoppers, a significance level of p < 0.00001 was revealed and C16:3-solamine was undetectable in individuals feeding on Solanum tuberosum ‘Gala’ ( Figure 5 ). In intestines of planthoppers, a significant difference of this compound was observed with a p-value of 0.001. Differentiation by the sex of the planthoppers demonstrated a significant uptake of the C16:3 solamine at p < 0.001 of the male and female planthoppers fed on S. bulbocastanum , thus no sex-specific differences in the feeding behavior of plant hoppers were observed ( Figure 6 ). Download figure Open in new tab Fig. 5. Boxplots of the C16:3-solamine peak area in extracts of whole planthopper bodies (A) and extracts of intestines (B) of planthoppers after feeding on Solanum bulbocastanum vs. Solanum tuberosum cv ‘Gala’. The x-axis indicates the two plant species (here, Sblb2G: S. bulbocastanum , Gala: S. tuberosum cv ‘Gala’), the y-axis indicates the Log10 peak area of the LC/MS data Download figure Open in new tab Fig. 6. Boxplots of the C16:3-solamine peak area in extracts of intestines of female and male planthoppers after feeding on Solanum bulbocastanum vs. Solanum tuberosum cv ‘Gala’. The x-axis indicates the two sexes (female P. leporinus , male P. leporinus ) fed on the two plant species (here, blb2G: S. bulbocastanum , Gala: S. tuberosum cv ‘Gala’), the y-axis indicates the Log10 peak area of the LCMS data of the C16:3-solamine 3.5 GC/MS analysis In total, 80 compounds of the collected VOCs of S. bulbocastanum and of S. tuberosum cv ‘Gala’ resulted after selection as described in the section ‘Material and Methods’. A table with all compounds is provided in Online Resource 1 (ESM_1.pdf). Thereof, 32 compounds were found in ‘Gala’ only and seven compounds only in the wild potato. These differentiating compounds are listed in Table 4 . Chromatograms of the GC/MS data of the collected VOCs show the very distinct profiles of the two species ( Figure 7 ). Compounds which contribute significantly (p<0.001) to a differentiation of the VOC patterns are inserted as arrows in a non-metric multidimensional scaling plot ( Figure 8 ). View this table: View inline View popup Table 4. List of identified, annotated or unknown volatile organic compounds only found in S. bulbocastanum or S. tuberosum cv ‘Gala’, indicating the retention indices (RI), p-values and species. Compounds differentiating the VOC pattern after application of environmental vector analysis with a permutation of 10,000 are listed with following significance codes: ‘***’ for p < 0.001, ‘**’ for p < 0.01, ‘*’ for p < 0.05 Download figure Open in new tab Fig. 7. Total ion current chromatograms of volatile organic compounds of S. tuberosum cv ‘Gala’ (A) and S. bulbocastanum (B) with arrows pointing to selected compounds; dodecanoic acid, ethyl ester was used as internal standard (IS) Download figure Open in new tab Fig. 8. Non-metric multidimensional scaling (NMDS) plot classifying the VOC profiles of S. bulbocastanum and S. tuberosum cv ‘Gala’ based on Bray-Curtis dissimilarities, displayed as triangles and asterisks, respectively. Compounds (c i ) accounting for a differentiation of the VOC pattern (with p-values < 0.001) after vector analysis are displayed by arrows. A numbered list of compounds is provided in Online Resource 1 (ESM_1.pdf). The significance of fitted vectors is assessed using permutation (level 10,000) of the 80 target compounds, the goodness of fit statistic is squared correlation coefficient (r 2 ) As shown by the results of the multivariate statistical test PERMANOVA ( Table 5 ), the difference of the two species clearly affects the VOC profiles (with a p-value of 0.0019). It accounts for 70.4 % (R 2 = 0.704) of the variation in the composition. The PERMDISP test results in a p-value of 0.7133, thus showing no evidence of significant difference in dispersion between the two species. This agrees with the visualization in Figure 8 . View this table: View inline View popup Download powerpoint Table 5. Results of multivariate statistical tests PERMANOVA and PERMSDISP based on Bray-Curtis dissimilarities for the compositional data set. The number of permutations was 10,000 Discussion For the first time plant-insect-interactions between the vector P. leporinus and the wild potato S. bulbocastanum have been investigated. Mortality experiments designed as no-choice experiments revealed lower survival rates of the vector compared to survival rates on a common potato variety. Since these first results seemed astonishing, subsequent experiments followed, including dual-choice-experiments to investigate whether S. bulbocastanum either elicits attraction or rather evasive behavior of the planthopper. The choice-experiments revealed a preference of the planthoppers towards the wild potato rather than the potato cultivar. A study of Gorzolka et al. (2025) discovered a long chain fatty acyl (LCFA) solamine (C16:3(7 Z ,10 Z ,13 Z )-solamine), which was isolated from the leaves of S. bulbocastanum . Its role in plant defense was investigated with interesting results on common potato pests such as bioactivity against the potato flea beetle and the Colorado potato beetle (CPB). Unlike P. leporinus , the CPB is not a plant sap sucking insect but feeds on leaf tissue. Likewise, the potato flea beetle feeds on leaf tissue pitting little holes into potato leaves. In feeding no-choice assays, CPB fed significantly less leaf material of Solanum bulbocastanum compared to the Solanum tuberosum varieties ‘Maxi’ and ‘Quarta’. Small quantities of exogenous applied C16:3-solamine led to significant lower feeding activity with > 70% less leaf consumption for the CPB and > 80% for the potato flea beetle. However, S. bulbocastanum leaf tissue or the exogenous applied C16:3-solamine were not sufficient to lead to acute toxic effects on these pests ( Gorzolka et al. 2025 ). Compared to the no-choice test of our study, we did not study the feeding activity of the planthoppers but their survival rate. However, the planthoppers sucked on both, the stalks of the wild potato and of the potato cultivar ‘Gala’ during these experiments. In downstream feeding experiments, blue stained plant sap was observed in both S. bulbocastanum and S. tuberosum treated planthopper intestines. This suggests that the planthoppers consumed xylem sap as the plant stalks were cut at the bottom and the top to allow for a quick water transport via transpiration which can be considered to take place in xylem cells ( Holbrook and Zwieniecki 2005 ; Kim et al. 2014 ). This latter finding adds new insights in the biology of this vector suggesting a flexibility in its feeding strategies which have not been studied so far. At the current status of knowledge, this would be rather non-characteristic for a vector of phloem-limited phytoplasma ( Weintraub and Beanland 2006 ; Gross et al. 2022 ), but has been observed before in another phloem feeding phytoplasma vector, the psyllid Cacopsylla picta ( Görg and Gross 2021 ). Thus, the feeding behavior of P. leporinus should be studied in more detail in future feeding experiments. Most of the planthoppers exposed to S. bulbocastanum were dead after a few days of exposure, i.e. about 68% in the first experiment after three days compared to 11% of the control, and about 83% in the second no-choice experiment after two days, compared to 43% of the control. In both experiments, more than 90% of the insects, which fed on S. bulbocastanum , were dead after four days. Since the feeding activity was not investigated at individual insect level and in absolute amounts, we can neither conclude nor exclude that this plant has deterrent effects on planthopper feeding behavior. However, choice experiments demonstrated that significantly more insects were observed on the wild potato than on the cultivated potato variety over a two-day period. Other studies would be necessary to investigate which uptake times would be sufficient for toxic effects. Moreover, feeding studies with isolated components such as C16:3-solamine or with plant extracts would be required to infer lethal effects of specific S. bulbocastanum compounds on the planthopper. To our knowledge, P. leporinus has not been studied yet in terms of behavioral studies, neither exist reports of artificial feeding systems with this species. That would be a pre-requisite for further investigation on toxic effects of acyl solamines in the planthopper. A proof of the uptake of the plant sap was provided by LC/MS analysis of whole planthoppers and their intestines. We have undertaken this analysis to see whether an ingestion of specific plant ingredients occurred, as those compounds might account for the lower survival rate of the planthoppers on S. bulbocastanum . It turned out that C16:3-solamine played a major role in differentiating the groups of planthoppers fed on S. bulbocastanum compared to those fed on S. tuberosum . In another study, ultrasonication-assisted methanol extraction of dried S. bulbocastanum leaves proved to be cytotoxic towards different breast cancer cell lines which was inferred to the phenolic profile of the extract revealing high levels of phenolic acids ( Paraschiv et al. 2022 ). Phenolic acid derivatives were also detected in our LC/MS analysis, but were not causing a significant differentiation of the profiles. With regards to possible olfactory cues of the wild species leading to attraction of the planthoppers, the VOCs of both species, S. bulbocastanum and S. tuberosum cv ‘Gala’ have been collected and analyzed. As the planthopper is directed towards the wild crop for at least a first preference, visual or olfactory cues or a combination thereof are likely to play a role in plant detection, as has been reported for other phloem sucking hemipteran insects ( Todd et al. 1990 ; Patt and Sétamou 2007 ; Wenninger et al. 2009 ; Aráoz et al. 2019 ; Gross et al. 2022 ). In our study, changes in growth stage, plant nutritional status and abiotic factors of the VOC emitting plants were not compared, i.e. it is unclear whether variations of such factors would have resulted in similar results of the VOC profiles or behavioral responses in the choice-experiments. Out of the 80 detected compounds, about half of them (32 + 7) are separating the VOC profiles of the two species. Given this, antennal responses of P. leporinus to compounds of these profiles could be further investigated by electroantennography. Subsequent behavioral tests with specific compounds could elicit potential vector behavior to attractant or repellent acting compounds. Resulting plant infections not only cause damage in yield and quality but also cause concern in potato growing regions, particularly those for seed production and for the food chain. This concern is justified given the reduction of tuber quality in terms of decreased germination capacity and a complete loss of the tradable commodity, but also with regards to critically increased contents of sugars leading to non-processable tubers for the food industry and reduced storage capability ( Mahillon et al. 2025 ; Lang et al. 2025 ). Consequently, emergency authorizations of insecticides have been approved in Germany in the growing season 2025 for application in potato and sugar beet for the control of P. leporinus , but do neither allow for a long-term solution to this pest, nor to the bacterial pathogens transmitted by it. Given the wide range of host and feeding plants besides potato and sugar beet, this vector poses major challenges for agriculture including the current cropping rotation. Studies of the cropping rotation as a potential to reduce the vector population of P. leporinus are sparse ( Bressan 2009 ; Pfitzer et al. 2024 ). Besides, these studies were conducted before the vector expanded its feeding and host range. Neither the use of repellent crops or substances nor the use of trap crops have been reported so far for P. leporinus . Our results suggest that S. bulbocastanum is more attractive to P. leporinus compared to conventional potato varieties, and at the same time exhibits lethal effects. Given this, S. bulbocastanum might serve to attract the vector, and, doing so, reduce it as well as the pathogen population which may decrease because of a lower infection density in field crops. However, this observation lacks proof at field level or at least larger scale, i.e. in field plots. In this scenario, S. bulbocastanum might serve as a trap plant, an already existing natural variant of the principle of “attract and kill” ( Pickett and Khan 2016 ; Gregg et al. 2018 ). With regards to trap plants, Killiny et al. (2021) investigated the effects of biophytoene desaturase-silenced citrus to lure Diaphorina citri , the phloem sucking vector of the Huonglongbing disease. Silenced plants expressed visual, olfactory, and gustatory cues and led to a higher attraction of the psyllid vector compared to other citrus plants of the orchard ( Killiny et al. 2021 ). Chaste tree ( Vitex agnus-castus ) was studied as a trap plant in vineyards for Hyalesthes obsoletus [Hemiptera: Cixiidae], a planthopper of the same family as P. leporinus and also a vector of the phytoplasma ‘ Ca . Phytoplasma solani’ in grapevine and potato ( Maixner 1994 ; Ulrich et al. 2010 ). Finally, chaste tree was not recommended as it takes up the transmitted pathogens and acts as a source of infection itself ( Moussa et al. 2019 ). With regards to trap plants and traps in general, there is a knowledge gap for P. leporinus . The vector could be attracted at adult stage aboveground but also belowground at nymphal stage according to its lifecycle ( Behrmann et al. 2022 ). With regards to the economic impacts and the need for infection controls, it would be a very interesting question, if S. bulbocastanum itself could be infected by ‘ Ca . Phytoplasma solani’ and ‘ Ca . Arsenophonus phytopathogenicus’, and if bacteria could proliferate in S. bulbocastanum and if symptoms would occur. Since this planthopper not only expanded its feeding and host plants, but also its spatial distribution, various levers are expected to be used in its future management. Tolerant and resilient crops as well as biotechnological plant protection applications that make use of manipulating vector behavior or interfering vector-plant-interactions would form part of such a strategy. Also, commercially available potato cultivars already possessing genes from S. bulbocastanum could be screened for increased pathogen tolerance or effects on the vector. Conclusion To our knowledge, results of P. leporinus and its interaction with a wild potato, S. bulbocastanum , are presented for the first time by this study. These results could be a starting point for further research directed towards natural plant defense and sustainable crop protection such as potato resistance breeding. S. bulbocastanum plants, the application of its leaf extracts or volatile organic compounds could be further investigated for attract- and-kill strategies of the vector P. leporinus . Their potential as plants for intercropping between potato ridges or as plants bordering potato fields could be investigated in field trials, too. Funding The main part of this study was implemented within the project SIKAZIKA, funded by the European Union and the State Hesse, Germany within the framework of the European innovation partnership for agricultural productivity and sustainability (EIP-Agri, grant number 9100443-INVEST-1). Acyl solamine and Solanum bulbocastanum LC-MS analyses were funded by the Federal Ministry of Agriculture, Food and Regional Identity (BMLEH) based on a decision of the Parliament of the Federal Republic of Germany via the Federal Office for Agriculture and Food (BLE) [grant number 28A8706A-C19]. Notes The authors declare no conflicted or financial interests. Data Availability The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request. Author Contribution Statement ET, JG and KG conceived and designed research. ET and KG conducted experiments. ET and KG analyzed data. ET wrote the manuscript to which KG contributed sections related to LC/MS analysis. JG, KG and ET revised the manuscript. All authors read and approved the final manuscript. Acknowledgements We thank David Löffler and Sanela Kugler for their support in catching planthoppers for the trials, as well as the TH Bingen, Germany for allowing us to collect planthoppers on their fields. Furthermore, we are grateful to Sanela Kugler and Svenja Stein for their excellent assistance with the experiments, Ali Karimi and Alicia Koßmann for their assistance in setting up the GC/MS analysis, and Jannicke Gallinger, Dominik Schmidt and Andreas Jürgens for their support in statistical questions. 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