Motivational trade-offs in a cockroach: impact of conditions, injury, and development

Motivational trade-offs in a cockroach: impact of conditions, injury, and development | 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 Confirmatory Results Motivational trade-offs in a cockroach: impact of conditions, injury, and development Joe Jenkinson , Joanna Logan , Christopher Robertson , Keith Lockhart , View ORCID Profile David N. Fisher doi: https://doi.org/10.1101/2025.10.31.685749 Joe Jenkinson 1 School of Biological Sciences, University of Aberdeen, King’s college , Aberdeen, AB243FX, United Kingdom Find this author on Google Scholar Find this author on PubMed Search for this author on this site Joanna Logan 1 School of Biological Sciences, University of Aberdeen, King’s college , Aberdeen, AB243FX, United Kingdom Find this author on Google Scholar Find this author on PubMed Search for this author on this site Christopher Robertson 1 School of Biological Sciences, University of Aberdeen, King’s college , Aberdeen, AB243FX, United Kingdom Find this author on Google Scholar Find this author on PubMed Search for this author on this site Keith Lockhart 1 School of Biological Sciences, University of Aberdeen, King’s college , Aberdeen, AB243FX, United Kingdom Find this author on Google Scholar Find this author on PubMed Search for this author on this site David N. Fisher 1 School of Biological Sciences, University of Aberdeen, King’s college , Aberdeen, AB243FX, United Kingdom Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for David N. Fisher For correspondence: david.fisher{at}abdn.ac.uk Abstract Full Text Info/History Metrics Data/Code Preview PDF Abstract The ability to flexibly trade off access between competing or antagonistic stimuli (motivational trade-offs) is a key criterion for assessing whether animals can feel pain. However, in some commonly farmed, exterminated, and studied insect orders such as Blattodea (cockroaches and termites), the ability to make motivational trade-offs is unconfirmed, which makes it difficult to assess their welfare needs. Here we gave cockroaches a choice between access to shelter or nutrition across a series of linked experiments to assess how motivational trade-offs are flexibly adjusted due to external conditions and injury and develop across ontogeny. We found cockroaches would adjust how willing they were to access nutrition instead of shelter based on the intensity of the light they would be exposed to and how long they had been deprived of food. They can therefore make flexible motivational trade-offs. We also found that injury reduces willingness to accept light exposure when accessing nutrition, which is consistent with the idea of the sensation of pain reducing risk-taking behaviour. Finally, we showed that juvenile cockroaches can make motivational trade-offs, but only towards the end of their development, and that young juveniles do not trade-off. All together our results provide comprehensive evidence that cockroaches can make highly flexible motivational trade-offs, filling in a gap in our understanding of cognitive abilities in insects linked to their ability to feel pain. These findings indicate the need to design legislation, housing, and production methods that account for and enhance invertebrate welfare. Introduction Animal welfare has varied definitions but typically refers to the quality of life experienced by the animal, where negative physical and psychological states are minimised and positive ones promoted ( Hewson, 2003 ; Reimert et al., 2023 ). Considerations include suitable nutrition, space to move, freedom from fear, pain and disease, and the allowance of natural behaviours ( Mellor et al., 2020 ). Understanding and improving welfare is relevant not only in agricultural settings, but also in scientific research and for wild animals. Researchers have a duty of care to minimise animal suffering when using animals for experimentation or other practical purposes ( Drinkwater et al., 2019 ), and in wild animals, as welfare can be an important component of conservation ( Paquet & Darimont, 2010 ). In order to be protected under animal welfare legislations the animal must be deemed sentient and so have the capacity for suffering. Key in concluding whether an animal can suffer is whether it feels pain; an unpleasant sensory and emotional experience associated with, or resembling that associated with, actual or potential tissue damage ( Raja et al., 2020 ). There is a broad consensus that vertebrates including birds, amphibians, fish and mammals feel pain ( Gentle et al., 1991 ; Machin, 1999 ; Sneddon, 2003 ; Williams, 2019 ), supported by behavioural indicators such as protective responses, changes in motivation, and avoidance learning following injury ( Rutherford, 2002 ). In addition, these pain-feeling animals possess physiological indicators for the ability to feel pain such as nociceptors, complex neurological pathways linking sensory brain regions, and modulation of these nociceptive pathways when exposed to analgesics ( Crump et al., 2023 ; Gibbons, Sarlak, et al., 2022 ; Key et al., 2021 ). However, invertebrates (approx. 97% of animal species) such as insects have traditionally been considered to lack key physiological indicators and not to show the behavioural indicators that indicate they feel pain ( Adamo, 2016 , 2019 ; Eisemann et al., 1984 ), and so most animal welfare legislation excludes invertebrates. For example, in the UK the only invertebrates protected under legislation during scientific research are cephalopods (Animals (Scientific Procedures) Act 1986, 2021). Feeling pain is supposed to help an animal avoid potentially damaging situations in the future, a putatively adaptive shift ( Bateson, 1991 ; Crook et al., 2014 ). The supposed adaptive value of pain suggests that many taxonomic groups may possess pain-sensing mechanisms, not solely vertebrates. Recently, the evidence for behavioural and physiological indicators of the ability to feel pain has been re-assessed for decapod crustaceans and cephalopod molluscs ( Birch et al., 2021 ) and for insects (Gibbons, Crump, et al., 2022 ). These reviews consistently found that, when we have looked for the presence of the physiological and behavioural indicators in these taxa (often we have not), we find them, strongly suggesting they may feel pain. However, gaps in evidence remain, limiting our ability to be decisive about whether invertebrates can suffer and so if their welfare requires formal legal protection. The behavioural indicators of pain are each more than a mere reflex and demonstrate some longer-term shift to response to a noxious stimulus ( Bateson, 1991 ; Sneddon et al., 2014 ). For example, motivational trade-offs require an animal to demonstrate a flexible trade-off between competing stimuli, such as accepting exposure to a noxious stimulus to gain a reward or avoid an even more noxious alternative. Motivational trade-offs are thought to stem from adaptive decision making, where animals in the wild must weigh up several competing factors against each other e.g., shelter availability vs predation risk ( Lima & Dill, 1990 ). The presence of motivational trade-offs across different domains, showing the processing of different types of information into a common currency, indicates central processing of stimuli ( Crump et al., 2022 ), which is important for inferring more complex cognition. Examples of motivational trade-offs in invertebrates include shore crabs ( Carcinus maenus ), that trade-off exposure to electric shocks against exposure to bright light ( Barr & Elwood, 2024 ), and Drosophila melanogaster , which show some willingness to tolerate 120V shocks to gain a reward of ethanol but not to gain the less-rewarding sucrose ( Kaun et al., 2011 ). However, in insects motivational trade-off research has focused on Diptera and Hymenoptera (representing approx. 27% of insect species), with a knowledge gap across other orders (Gibbons, Crump, et al., 2022 ). One well-studied insect order without evidence for motivational trade-offs is Blattodea (cockroaches and termites). Blattodea are exceptionally widespread, featuring in nearly all climates with several species being well-known as domiciliary pests and disease vectors ( Bell et al., 2007 ). Cockroaches are also widely used in neuroscience research ( Huber et al., 1990 ). Despite these factors, there are no studies testing whether cockroaches make motivational trade-offs (although Varnon and Adams (2021) did find that the presence of food inhibited startle responses to light in the orange head cockroaches ( Eublaberus posticus ), suggesting the ability to integrate information across domains if not evidence for a trade-off between accessing competing goals). We aim to fill the gap around motivational trade-offs in Blattodea here. Additionally, we explored motivational trade-offs in cockroach nymphs, as Gibbons et al . (2022) highlighted that no juvenile insects have been tested for the ability to make motivational trade-offs. It is crucial that juveniles are tested, as many insects spend the majority of their lives, especially in production settings, as juveniles, and so their capacity to suffer in this life stage is key. We carried out three linked experiments to assess the extent of motivational trade-offs in adult and juvenile cockroaches. We used a trade-off between access to nutrition (a positive stimulus) vs. exposure to light (a negative stimulus) to test for motivational trade-offs. First, we tested whether conditions altered the magnitude of the trade-off between access to nutrition and exposure to light. Demonstrating the flexibility of motivational trade-offs is key for understanding whether these are simple reflexes or evidence of central processing. Second, we determined if an injury altered this trade-off. Injury altering subsequent behaviours that indicate an increase in protective behaviour is considered key for showing the feeling of pain rather than simply nociception ( Sneddon et al., 2014 ). Finally, we determined if juveniles made the trade-off, and if the magnitude of the trade-off varied with mass, which we used as a proxy of developmental stage ( Wu, 2013 ). As mentioned above, no juvenile insects have been assessed for their ability to make motivational trade-offs, yet this is a very important life stage, especially in production settings where many individuals may be harvested as juveniles ( Barrett & Fischer, 2023 ). We used the Blaberid cockroach Blaptica dubia , a species widely used in the pet trade as live feed but also increasingly used in scientific research. We predicted that cockroaches would show motivational trade-offs and that the magnitude of the trade-off would vary in response to conditions and be influenced by injury. For juveniles, we predicted they would make motivational trade-offs and further predicted that larger individuals would be more likely to perform trade-offs as the need for resources with larger moults is more pronounced than when smaller. While no one experiment is conclusive evidence of the ability to make motivational trade-offs, we aim to provide three lines of evidence, allowing us to arrive at a general conclusion on the probability that cockroaches are capable of motivational trade-offs ( Brown & Birch, 2025 ). Methods Stock and housing We used a population of B. dubia that have been maintained at the University of Aberdeen since 2021, with periodic purchase of new individuals to limit inbreeding. The B. dubia stock population are stored in an insectary room within 48L plastic (polypropylene) source boxes (60cmx40cmx35cm, Really Useful Boxes, Really Useful Products Ltd.) each containing several 24-egg sized cardboard carton for shelter, which are replaced after signs of degradation. Source boxes are cleaned twice weekly with 70% ethanol solution. Hydration in the form of three 40g carrot slices and food in the form of 10g of Sainsbury’s Complete Nutrition Adult Small Dog Dry Dog Food (rough nutritional values: 1527kJ energy, 24g protein, 12g fat per 100g) are also replaced twice per week. This insectary remained between 28-30°C under 50% humidity with a 50:50 light/dark light system that kept lights on only between 7am and 7pm (further details on husbandry are provided in Fisher, 2023 ). Base Method We conducted three different experiments using the same base method, with differing variables explored in each experiment to test for the existence of motivational trade-offs and their extent in B. dubia . In each experiment we tested for motivational trade-offs by comparing the willingness of individuals to stay in one side of a box that was empty, but sheltered, or go to the other side of the box that contained a piece of carrot, which was one of two treatments: either exposed to light (“exposed”) or sheltered (“sheltered”). By comparing these two groups we can determine how much the cockroach goes to the carrot depends on being exposed to or sheltered from light, indicating a trade-off between access to shelter and access to nutrition. Further, we can vary the experimental conditions or characteristics of individuals to determine the flexibility of the trade-off. Experiment 1 explored the impact of conditions (light intensity or hunger), experiment 2 the impact of injury, and experiment 3 the impact of development. We only used males for our experiments with the adults as females from the stock population were being used in other experiments, and it was easier to injure the males for experiment 2 due to their exposed wings. We did not sex the nymphs for the experiment on juveniles and so assume the experiment included an equal number of males and females. We filmed all trials with ABUS IP video surveillance 8MPx mini tube cameras (ABUS, Germany) through the “AnyCam” software (AnyCam.iO) rather than carry out direct observations, to limit the impact of observer presence on behaviour. In the first step of the base method, we randomly selected cockroaches from their 48L source boxes and placed them into a separate environment without access to nutrition or hydration for 24 hours. We did this to increase the cockroaches’ motivation to leave the always-sheltered side of the box and access the carrot. For experiment 1, we placed cockroaches separately into small boxes (7.9cm x 4.7cm x 2.2cm), while for experiments 2 and 3 individuals were held together in a 48L box. After the 24 hours we transferred each individual cockroach to a 0.9L box (22cm x 10cm x 7cm) that was either 50% or 100% shaded depending on whether they were in the exposed or sheltered treatment respectively ( Fig. 1 ). Shade was created using plain white A4 paper that was cut to shape and taped around the top and sides of the boxes excluding one side to allow lateral visibility into each box. In each box we placed a piece of carrot of approximately 3g. In experiment 1 we added carrots to the boxes before the cockroaches were taken to the recording room, whereas for experiments 2 and 3 we added their carrot slices at the point filming began. These 0.9L boxes were cleaned with 70% ethanol between trials to limit any effect of scent or contaminants on the next cockroach to enter the box. Download figure Open in new tab Figure 1. Lateral (left) and dorsal (right) views of the (A) Exposed box and (B) Sheltered box (filming is conducted from top-down). We transferred cockroaches in individual 0.9L boxes to the recording room, with an equal mix of exposed and sheltered treatments. We heated the recording room with portable heaters to approximately 25°C for at least an hour before beginning any trials, while we recorded temperature during trials with a thermometer and controlled for its effect statistically (see Data analysis). As the recording room was not sealed, to prevent cockroach escape into the wider building we placed each 0.9L trial box into a larger box for filming. For trial 1, we placed two randomly ordered pairs of 0.9L boxes in larger 48L boxes. We placed a single camera per 48L box (covering two 0.9L boxes) 6.6 cm above the ground at a distance of 30cm from the 0.9L boxes ( Fig. 2A&B ) and used up to four cameras simultaneously to assess up to eight individuals at once. For trials 2 and 3 we placed the 0.9L boxes singly into 5L boxes for recording, using one camera (elevated by 14cm) per four boxes by filming two side-by-side and stacking an extra pair of empty 5L boxes behind the first two to prop up the next two ( Figure 2C&D ). We used up to two cameras simultaneously allowing us to assess up to eight individuals at once. We positioned these cameras to provide a clear side view of all the boxes in front so that all cockroaches and their position were always clearly visible in recordings, and since the conditions for the cockroaches do not change between the set-ups (they all experience a 0.9L box with a piece of carrot and either exposed or sheltered treatment) we believe the experiments to be comparable. We randomly assigned cockroaches to the camera and location in the arrangement we placed them in. We placed a pet heat mat (models TK-HPP7040A & TK-HPP6540, OnKey Electronic Technology Co. Ltd) underneath each of the paired 5L or individual 48L boxes containing cockroaches ( Fig. 2 ) to maintain them at a recommended temperature of ∼30 ° C so that low temperatures did not influence their locomotive ability ( Alamer & Hoffmann, 2014 ). Download figure Open in new tab Figure 2. Diagram of experiment set up for experiment 1 lateral (A) and dorsal (B) views and experiments 2 and 3 lateral (C) and dorsal (D) views. Each set up was designed to give a clear view of the open side of each 0.9L box so that the position of the cockroach within could be determined. We set the cameras to record for 30 minutes and then immediately left the recording room. At the end of experiment 1 we transferred cockroaches back to the source box they came from. Cockroaches could thus have been used more than once between different trial days but would not be used in the same trial day or consecutive days due to being preselected for isolation the day prior. This provided time for readjustment to their source box environments after trials before potentially being used again on another trial day, negating any behavioural impacts. In experiment 2 all cockroaches were frozen in -20°C after the experiment, which is thought to be a quick and so humane way to kill this species ( Tucker et al., 2023 ), and so were never re-used. In experiment 3 nymphs were returned to the stock population but were not re-used in the experiment. Experimental Conditions In experiment 1 we ran three separate trials, all underpinned with the base method. The first trial was only the base method. In the second trial we then tested the impact of conditions by altering the light level in the open half of the exposed boxes by attaching translucent paper to partially block light over the food as opposed to allowing full light. Third, we explored the effect of changing the starvation period from 24 hours to 48 hours in the final trial of experiment 1. The experimentation period (including laboratory testing, video extraction, and filming) ran from 11/10/2022 to 22/11/2022 and used 72 individuals, although recordings were lost for 11 of these, leaving 61. For experiment 2 we used the base method, but after the 24-hour starvation period all the starved cockroaches were randomly allocated as injured or uninjured. For the injured individuals, we cut off 1cm of their outer wings with sharp scissors. The wings were clipped as B. dubia wings in males have limited function and so wouldn’t impact locomotion for the experiment. We handled uninjured cockroaches for around 40 seconds, which is the approximate time the injured cockroaches spent being handled while having their wings clipped. Experimentation ran from 21/10/2024 to 12/11/24 and used 120 individuals, but data from 16 trials were lost due to camera failure, leaving 104. For experiment 3 we used body mass as a proxy for age/developmental stage. We weighed each nymph to be used in the experiment before the starvation period using a Fisherbrand Analytical Balances, readability 0.0001g scales. Body mass was recorded to two decimal places. Experimentation ran from 28/10/2024 to 20/11/24 and used 80 individuals. Video and data analysis For each experiment we followed a similar analysis approach. We first removed the first 5 minutes of each trial as during this “habituation” period the cockroach may not have been responding to the experimental treatments but rather to the novelty of the box. We watched the remaining video and every 60 seconds (experiment 1) or every 30 seconds (experiments 2 and 3) counted the number of times during the trial the cockroach was either on the same side of the box as the carrot, the opposite side, in the middle, its side could not be determined (experiments 2 and 3 only), and the number of times it could not be seen (experiment 3 only). We increased the sampling rate for experiments 2 and 3 to give a higher resolution on behaviour. For experiments 1 and 2 there were no time points where the cockroach could not be seen, and so both of these were analysed in the same way. For experiment 1 analysed data from all three conditions (base method, reduced light, longer starvation) together. We fitted a Poisson GLM with number of time points the cockroach was on the same side as the carrot as the response variable, and predictor variables of exposed/sheltered as a two-level factor, conditions (full light/reduced light/starvation) as a three-level factor, the interaction between these two terms, and temperature (scaled to a mean of 0 and a variance of 1, which aids the fitting and interpretation of regression models; Schielzeth, 2010 ) as a continuous covariate. For experiment 2 we fitted a Poisson GLM with the same response variable, with predictor variables of exposed/sheltered as a two-level factor, injured/uninjured as a two-level factor, the interaction between these two terms, and temperature (again scaled to a mean of 0 and a variance of 1) as a continuous covariate. For experiment 3 there were eight trials where the cockroach could not be seen at four time points, and so we needed to account for a different number of overall observations. We therefore fitted a binomial GLM with number of times the cockroach was the same side as the carrot as the “successes” and the total number of observations where the individual could be seen as the number of “trials”, which accounts for variation among cockroaches in the latter value. We included predictor variables of exposed/sheltered as a two-level factor, mass in grams as a continuous variable (also scaled to a mean of 0 and a variance of 1), the interaction between these two terms, and temperature (again scaled to a mean of 0 and a variance of 1) as a continuous covariate. For all models we calculated p values with a type III sum of squares ANOVA in the package car ( Fox, 2016 ). The main effect of exposed/sheltered indicates whether the cockroach is more or less willing to go to the carrot when it would be exposed to light in doing so, providing evidence of a motivational trade-off. Meanwhile the interaction between exposed/sheltered and the other factor in each analysis indicates whether this trade-off depends on conditions (experiment 1), injury (experiment 2), or development phase (experiment 3). Ethical note While no formal ethical approval was required for this work, we adhered to ASAB guidelines throughout. Our approach was minimally invasive aside from in experiment 2 where we were required to injure the cockroaches to test our hypothesis. These individuals were tested immediately after injury and euthanised directly after the experiment to minimise any period of potential suffering. We also deprived the cockroaches of food and hydration for 24 or 48 hours, but this is well within conditions cockroaches are naturally exposed to and are specifically adapted for ( Bell et al., 2007 ). At the end of each study animals were returned to the stock population unless stated otherwise. Results Experiment 1 (impact of conditions) Cockroaches avoided the carrot more when it was exposed to light (main effect of exposed/sheltered, incidence rate ratio [IRR] = 0.640, χ 2 1 = 28.878, p < 0.001), indicating a motivational trade-off. Additionally, the magnitude of this trade-off varied among conditions (interaction between exposed/sheltered and condition, χ 2 2 = 6.597, p = 0.037); the difference between exposed/sheltered was greater when the cockroaches were starved than in the full light condition (IRR = 0.590) and reduced when the intensity of light was reduced (IRR = 0.770; Fig. 3 ). The average preference for carrot also differed among conditions (main effect of conditions, χ 2 2 = 10.527, p = 0.005, IRR for starvation = 1.168, IRR for reduced light = 0.947). Cockroaches showed reduced preference for carrot in warmer temperatures (β = -0.077, χ 2 1 = 12.151, p < 0.001). Download figure Open in new tab Figure 3. Dot and violin plots showing the cockroaches’ relative preference for carrot (number of times with carrot [max 25]) in exposed (blue) and sheltered (orange) arenas from experiment 1. We show the results for each experiment separately to illustrate how the differences between exposed and sheltered differ between the experiments (full light, food deprivation, and reduced light from left to right). Black lines show the medians, white lines the 25 th and 75 th percentiles. Experiment 2 (impact of injury) In experiment 2 cockroaches again showed a motivational trade-off between access to food and avoidance of light (main effect of exposed/sheltered, IRR = 0.842, χ 2 1 = 9.667, p = 0.002). The cockroaches made an even greater trade-off when they were injured (interaction between exposed/sheltered and injury, IRR = 0.636, χ 2 1 = 10.508, p = 0.001; Fig. 4 ). The average tendency to go to the carrot was also less when injured (min effect of injury, IRR = 0.840, χ 2 1 = 9.559, p = 0.002). Cockroaches showed reduced preference for carrot in warmer temperatures, although in this experiment not clearly so (β = -0.028, χ 2 1 = 1.772, p = 0.183). Download figure Open in new tab Figure 4. Dot and violin plots showing the cockroaches’ relative preference for carrot (number of times with carrot [max 50]) in exposed and sheltered arenas and when injured or uninjured from experiment 2. Dark blue shows exposed and uninjured, orange sheltered and uninjured, light blue exposed and injured, yellow sheltered and injured. Black lines show the medians, white lines the 25 th and 75 th percentiles. Experiment 3 (impact of development) Experiment 3 showed nymph cockroaches also make a motivational trade-off (main effect of exposed/sheltered, log-odds ratio = 0.727, χ 2 1 = 200.859, p < 0.001). The trade-off was minimal in smaller (younger) individuals and became more apparent in larger (older) ones (interaction between exposed/sheltered and mass, χ 2 1 = 35.515, p < 0.001; Fig. 5 ). Heavier individuals showed reduced preference for carrot (β = -0.630, χ 2 1 = 116.614, p < 0.001). Cockroaches showed less preference for carrot in warmer temperatures, although in this experiment not quite clearly so (β = -0.058, χ 2 1 = 3.287, p = 0.070). Download figure Open in new tab Figure 5. Scatterplot showing the cockroaches’ relative preference (number of times with carrot/total times observable) for carrot in exposed (blue) and sheltered (orange) arenas across sizes (developmental stages) from experiment 3. The difference between exposed/sheltered increases at higher masses. Trend line drawn in ggplot2 ( Wickham, 2009 ) using a Poisson GLM (with the proportion as a response variable, which differs from our analysis in the main text which used a Binomial GLM on counts of success and failures but should still give an accurate trend line). Discussion We found comprehensive evidence that B. dubia cockroaches can make motivational trade-offs. Cockroaches avoided accessing nutrition when doing so meant exposure to light, and the strength of the trade-off was affected by the strength of the light (less intense light meant relatively more willing to go to the nutrition) and the value of the nutrition (great need due to a longer dehydration period meant a smaller effect of light). When we injured cockroaches, they were less willing to go to the nutrition, and especially less willing to do so when the nutrition was exposed to the light, suggesting greater perceived risk and/or self-vulnerability. Finally, the trade-off between nutrition and shelter was present in large nymphs but less so or was absent in smaller nymphs, indicating the development of the ability to make the trade-off through ontogeny. Animals that react to negative stimuli through more than reflexes but with flexible behavioural adjustments that mitigate their risk of future threats are thought to be more likely to have the capacity to feel pain and therefore suffer ( Gibbons, Sarlak, et al., 2022 ; Sneddon et al., 2014 ). While simple trade-offs can be produced by cognitively simple animals ( Read & Nityananda, 2025 ) and trade-offs alone are not evidence for feeling pain ( Brown & Birch, 2025 ), along with other lines of evidence finding motivational trade-offs in any taxa increases the likelihood they feel pain. Demonstrating this landmark in insects and other invertebrates has therefore been considered a major development that will catalyse a change in how we consider the welfare of non-vertebrates, but evidence has been lacking in key insect orders such as Blattodea and for any juvenile insect ( Gibbons, Crump, et al., 2022 ). Our work is therefore a small but important piece of the puzzle of whether insects feel pain, which the collective weight of studies now firmly indicating they do. Although even bacteria might adjust behaviour in the presence of competing motivations ( Read & Nityananda, 2025 ), the clear interactive effects in each of our models show that the cockroaches are not simply producing a response that is the sum of the two stimuli but integrating different types of information to produce an appropriate response. Our first experiment showed that the degree to which cockroaches trade-off between access to shelter and nutrition is flexible with conditions. This flexibility is key for the subsequent inference that the trade-off is not merely a reflex and therefore evidences cognitive flexibility. Given adult cockroaches have previously been shown to meet six of the eight criteria of pain ( Gibbons, Crump, et al., 2022 ; with the other two simply not examined) it should therefore not be surprising that we were able to confirm one of the outstanding criteria: motivational trade-offs. We should note that the responses of the cockroaches in our first experiment were not exactly as predicted. For instance, when we reduced the intensity of the light, we predicted a smaller difference between exposed and sheltered, which is what we saw, but this came about through a decrease in willingness to go to the carrot when it was sheltered (the condition that had not changed) rather than increased willingness to go to the exposed carrot (the condition that had changed). Still, across all three studies we report we feel the evidence for motivational trade-offs is robust. The final frontier for pain in Blattodea is therefore whether they show preference for analgesia when injured (Gibbons, Crump, et al., 2022 ). Our second experiment found injured cockroaches were less willing to access nutrition, especially when that nutrition was exposed to light. That the trade-off between access to nutrition and exposure to light was more pronounced in injured cockroaches suggests that damaged cockroaches regard the environment as especially risky or themselves as especially vulnerable; what we would expect if feeling pain was adaptive ( Elwood, 2011 ). Note that since we injured cockroaches on their wings, which they do not use for locomotion, we can be confident that they did not access the nutrition less simply because their movement was impaired. American cockroaches ( Periplaneta americana ) with an abdominal puncture wound will groom more, and so that cockroaches could show flexible self-protection was known (see also: Gibbons et al., 2024 for similar wound tending behaviour in bumblebees Bombus terrestris ). We have taken this further by showing longer-term changes in behaviour that would appear to reduce the risk of further injury by exposing the animal ( Sneddon et al., 2014 ). Our finding has important implications for insect housing conditions – keeping certain species at high densities often leads to wing damage ( Wehmann et al., 2022 ), which we have shown here could be painful. Further, “procedures” on cockroaches in scientific experiments, such as ablating antennae or cutting holes in exoskeletons, may be painful, and as such we have a moral duty to minimise that pain when experimenting on animals for our own gain, such as by using appropriate analgesics ( Drinkwater et al., 2019 ). In our final experiment we showed that B. dubia nymphs also make a trade-off between access to nutrition and exposure to light. This is key as research on pain in juvenile insects is minimal compared to that in adults; Gibbons et al . (2022) identified no studies on motivational trade-offs in juvenile insects, and far less research on all eight criteria in juveniles in general. Given many insects spend most of their life as juveniles, before a short period reproducing as adults, and that in some production settings (e.g., for mealworm, juveniles of Tenebrio molitor beetles), the vast majority of individuals never reach adulthood before being slaughtered, demonstrating criteria for pain in preadulthood is especially important. What is particularly interesting about our result is how the magnitude of the trade-off changed with mass (which we used as a proxy for developmental stage). While larger individuals made a clear trade-off, smaller individuals did not and showed much less aversion to light. This could suggest that the ability to make the trade-off, the ability to sense the light, or the perception of risk posed by the light develop across ontogeny. It is not clear why the smallest juveniles would perceive the light as less risky, but cockroach eyes do grow between instars through the addition of new ommatidia ( Nowel & Shelton, 1980 ). To determine whether it is decreased motivation or decreased light sensitivity driving the lack of trade-off in small juveniles, we could assess light sensitivity across development, with a lower sensitivity at early stages suggesting reduced sensitivity as a cause. Alternatively, we could use different noxious stimuli, such as electric shocks used to assess motivational trade-offs in crustaceans ( Barr & Elwood, 2024 ; Elwood & Adams, 2015 ). Further work on juveniles in a wider range of species is also required to assess the general capacity for juvenile insects to make motivational trade-offs and so potentially feel pain, especially in commonly farmed species such as mealworm ( T. molitor ) and black soldier fly larvae ( Hermetia illucens ). In summary, we found evidence across three separate experiments consistent with the idea that B. dubia cockroaches make motivational trade-offs. This is a more complex level of cognition that insects were initially not thought to be capable of, but now the emerging consensus indicates that insects show the behavioural and neurological complexity sufficient to feel pain. Animal welfare implications Providing evidence consistent with the idea that insects feel pain indicates that insects kept in captivity (for commercial, research, and domestic purposes) should have housing conditions that minimise negative welfare and maximise positive welfare. Scientific research involving insects should also be subject to ethical approval as research on vertebrates (and cephalopods) is, although the degree may not be as extensive, and methods should be refined to minimise any suffering during procedures ( Crump et al., 2023 ; Drinkwater et al., 2019 ). Pest control methods can also be designed to minimise suffering through quick and painless deaths. The rapidly growing insects as food and feed industry will also need to consider the welfare of their “minilivestock” during housing, transport, and slaughter (Klobučar & Fisher, 2023 ; e.g. Rowe et al., 2024 for farmed crickets). Finally, we urgently need the development of better guidelines and new legislation to reflect evolving scientific knowledge on the capacity of insects to suffer (e.g. Fischer et al., 2024 ). Footnotes https://figshare.com/s/4f7ad21e2dcf33186389 References ↵ Adamo , S. A. ( 2016 ). Do insects feel pain? 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