Biomechanics Locomotion and Predation Escape in Cricket Species

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Longwood Senior Thesis Proposal O8ce of Student Research

Spring 4-17-2023

Biomechanics Locomotion and Predation Escape in Cricket Biomechanics Locomotion and Predation Escape in Cricket

Species Species

Kira M. Edens

Longwood University

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Biomechanical Locomotion and Predation Escape in Cricket Species

Applicant: Kira M. Edens ‘24

Farmville, VA 23901

Phone: 804-920-7309

Email: [email protected]

Faculty Mentor: Brandon E. Jackson, Ph.D.

Chichester Science Center, Room 305a

Department of Biological and Environmental Sciences

Longwood University

Farmville, VA 23909

Phone: 434-395-4836

Email: [email protected]

Discipline: Biology: Integrative Biology

Date of Submission: April 17, 2023

__________________________________ ___________

Kira M. Edens (Applicant) Date

__________________________________ ___________

Brandon E. Jackson, Ph.D. (Faculty Sponsor) Date

4/17/2023

Biomechanical Locomotion and Predation Escape in Cricket Species

I. Statement of Guiding Hypotheses

Locomotion is a fundamental to everyday life and ultimate survival of almost all living

animals through its role in capturing food, escaping predators, defense, finding a mate, and

discovering new habitats. Researchers have studied all different components of locomotion, from

flying to swimming and from running to jumping. These studies range drastically in numerous

organism species, specifically jumping performance in vertebrate and invertebrate animals and

insects. In the two semesters I have worked in Dr. Jackson’s lab, we have been surprised that

researchers have studied very little about the biomechanics behind a crickets jump. The few

studies worked with only one species of crickets to analyze the acceleration, power, velocity, and

work put out by the jumping performance. We have not found a single comparative study of

jumping techniques and mechanics among multiple cricket species which would show how their

morphology and physiology differ between jumps. Very few studies have been done on any

organism to examine how locomotor performance directly affects important behaviors like

escaping from predators.

This semester, Dr. Jackson and I set out a goal to film and track the locomotion of camel

crickets (Rhaphidophoridae Orthoptera) using the coding software Python and a tracking

software DeepLabCut. What we observed is that this cricket species has larger body and leg

lengths than other crickets. Cave Crickets (Rhaphidophoridae Orthoptera) have longer, more

slender legs than most other species (i.e. field crickets, Gryllinae Orthoptera). Biomechanical

theory says that longer legs should mean more time for force to be applied to the ground, which

can lead to higher energy and power exerted in a jump. Perhaps the different leg lengths could

aid them when jumping in predatorial escape compared to a shorter body and leg lengths. For

this Senior Thesis Research, I aim to investigate how longer body and leg lengths on crickets

jumping performance, both within and between two species of crickets, and how that jumping

performance affects actual predator escape. Thus, I hypothesis that cave crickets, with their

longer legs, will have a better jump performance than other species, and will have a higher

likelihood of escaping from predators.

II. Research Significance

There have been numerous prior studies involving crickets and insect jumps where

researchers observe different jumping and leg performance between multiple species from

humans to locust and jumping froghoppers to stinging jellyfish (e.g., Alexander, R, McN, 1995;

Patek, S. N., et al., 2022). Other researchers have specifically looked at ballistic movements of

jumping performance and the tasks of specialized appendages (e.g., Hustert, R. and Baldus, M.,

2010). Although there’s been little research pertaining jump performance of species between one

organism, crickets have specialized morphologies for jumping and are diverse, which would

make them an ideal model system for investigating how selected forces on jumping lead to the

diversity of extant crickets. Previous studies state that length of hind legs and body weight tend

to play a big role in locomotion of millimeter-to-centimeter sized insects, for the inertia force

that is depicted in jump performance is dependent on the weight and volume of the body (Li, F.

et al., 2012). The hind legs of an invertebrate major importance to jump performance, for when

hind legs extend and used as a propulsion for the muscles, generating a constant higher velocity

relative to the center of mass while producing low strain-rate that increases energy produced in a

jump (Alexander, R, McN, 1995)(Figure 2). In our lab we have already done preliminary data

that depicts the way crickets prepare for a jump. The crickets tend to move most of their body

when starting a jump performance where they tuck in their head and abdomen and tuck their hind

legs in to where the knees point outwards, so that the crickets create that power when extending

their hind legs (Figure 1). Studies have been done on female versus male body size of a

planthopper, and the females with a larger body size had a longer acceleration time and velocity

than males by a significant amount in both tests(Burrows, M., 2009).

When studying a close insect to crickets, researchers have discovered that bred and stress

induced grasshoppers have a takeoff velocity 1-2 times faster and 2-6 times longer jump

distances than wild grasshoppers due to morphological changes of matured grasshoppers and

unmatured grasshoppers (Hawlena, D., 2011). When observing crickets such as bush crickets

researchers found that female bush crickets’ hind leg femur length equivalates to four times that

of male bush crickets which contributed to a higher take-off velocity in females of

2.12±0.33ms

-1

while in males resulted in a lower take-off velocity of 1.51±0.2ms

-1

(Burrows, M.

and Morris, O., 2003). The extension of the hind legs is fascinating to researchers, for how much

height it brings to the center of mass to the cricket and how morphological differences in hind

leg length aid in jumping performance (Figure 2).

Researchers have also done many studies involving prey and predator interactions by

studying behavioral tactics producing in prey. For this study, we hope to determine how

differences in jumping performance is affected with body and length size of different cricket

species and predation escape (Figure 4). Previous studies of predation escape have shown that

crickets have sensor hairs that will alert them based on wind and a predator is close by,

researchers performed this by using airflow stimuli that could aid them in predatory escape

(Tauber, E and Camhi, J. M., 1995; Sato, N. et al. 2019). Our research and previous research

think that once stimuli happen, depending on morphological benefits such as longer legs would

aid the cricket in predatory escape, for longer hind legs result in a longer jump (Burrows, M. and

Picker, M. D., 2010)(Figure 1). For the second part of this research we aim to use lizards as a

predatory stimulant for the predatory-prey interactions. Research has been done on Varanid

lizards and their consumption rate of different species of prey, researchers elaborated that most

invertebrates and especially Orthoptera species are critical for a lizard diet due to habitation and

attraction (Cross, S. L. et al., 2020). This is important to our study, for the predation escape of

crickets will be more frequent and show differential data in the number of crickets remaining if a

predator that consumes invertebrates on a daily is used in this study. This project aims to be the

first study that we know of to directly attempt to link comparative cricket morphology, jumping

performance, and survival.

As described above, the findings of this project will be one of the first studies reported about

cricket species and the effect of body size and leg length have on jump performance and predator

escape. While our preliminary work on six camel crickets suggests that there is no effect of leg

length on jumping performance within the species, we do expect to find an effect when

comparing among species. In addition to other research done in the past on cricket jump

performance, we anticipate that other researchers could use this information to help determine

other insect or vertebrates jump performance between species – even if they are not studying

crickets.

III. Proposed Methodology

Animals and Animal Care

A collection camel crickets (Rhaphidophoridae Orthoptera), and field crickets (Gryllinae

Orthoptera) will be collected outside of the Environmental Education Center using a live insect

trap. Crickets will be housed in separate plastic containers depending on species and labeled.

Crickets will be provided access to food (fruits, vegetables, and grass), tap water, cardboard for

enrichment, air holes, and a humidity sensor.

A total of six mature unisex Anole lizards (Anolis carolinensis) (Carolina Biological

Supply Company) will be singly housed in singular 10 gallon (12”x12”x18”) glass terrariums

with potting soil as bedding. Lizards will be provided with food (Carolina Biological Supply

Company), plants and sticks for enrichment, tap water and misting daily, and a low-wattage

heating lamp to keep temperature at 24° C to 27° C with a warmer basking area that reaches

29.5° C to 33° C.

Jumping Performance Collection

A built glass arena lined with sandpaper will be used to hold crickets when filming

jumping performance. A high-speed camera (Edgertronic, Frame Rate: 500Frames/Second) and

LED lights was used to film crickets. Crickets will be individually placed in area and stimulated

to jump by air puffs from a disposable pipet to get about 10-20 jumps every week for three

hours. Once crickets are filmed with enough jumps they will be euthanized and preserved in 70%

ethanol.

Jumping Performance Tracking and Analysis

Each video will be tracked using DeepLabCut (Mathis, A. et al., 2018). The tracking

software will be trained using a video of previously filmed cricket videos by hand-labeling points

of the body: head, abrear (lower abdomen), hip, knee, and claw. The rest of the videos were

tracked using the trained tracking software (DeepLabCut). The preliminary filming and computer

model training has already been completed. The videos will then analyzed by the used of

calibration to provide femur length, tibia length, body length, jump start frame, jump end frame,

jump time, jump velocity, jump angle, jump height, jump power, mass specific jump power, and

relative mass. The data will be saved into an Excel file and then Excel was used to convert each

trial data from pixels to centimeters. Excel will be used to create acceleration, work, potential

energy, and kinetic energy from the data given.

Predator-Prey Interaction and Analysis

Crickets that will not be filmed for jumping performance will be used for predator-prey

interaction. A total of 24 crickets, four crickets per Anole lizard, will placed into each anole

lizard habitat container and left for 30 minutes. Two camel crickets (Rhaphidophoridae

Orthoptera) and two field crickets (Gryllinae Orthoptera) will be placed in each habitat. After

30 minutes, each habitat will be analyzed and recorded by counting how many crickets are still

alive to create a percent of crickets remaining (Figure 4). The crickets remaining were then

placed back into their normal habitat. This interaction was recorded another 4 times.

IV. Resources and Locations

a. Key Resources

i. Cricket housing facility, food, and water

ii. Lizard housing facility, food, and water

iii. Calibers

iv. Preservative fluids (70% ethanol)

v. Data analysis software

vi. High speed camera (Edgergonics)

vii. LED lights

b. Locations

i. All of the camera and recording resources required for this

experiment are located in Dr. Jackson’s research laboratory

Chichester 217

ii. Software programs such as Python, Excel, DeepLabCut, and

specialized computer to run the auto-tracking, are available in Dr.

Jackson’s Computer Image Analysis lab (Chichester 219)

V. Timeline for Completion

Summer 2022-Spring 2023

• Learn laboratory techniques/skills, including:

a. Digitizing video

b. Filming techniques

c. Programing (Python)

d. Built filming arena

e. Trained computer model to track cricket

• Collection of data for crickets

a. Filming

b. Statistical data analysis

c. Tracking via DeepLabCut and Python

Summer 2023

• Literature collection

• Analysis of preliminary data

Fall 2023

• Collection of cricket species at Environmental Education Center

• Build prey-predator filming arena

• Filming

a. Cricket species

• Tracking via DeepLabCut and Python

• Initial draft of LST paper, including analysis of preliminary data to Dr. Jackson by Spring

Break

• Presentation of preliminary data

Spring 2024

• Data Analysis

• Writing of Senior Thesis

• Final Senior Thesis submitted to Dr. Jackson

• Oral Defense Preparation

VI. Anticipated Committee Members

Brandon E. Jackson, Ph.D. Associate Professor of Biology, Longwood University

Charles D. Ross, Ph.D. Associate Professor of Physics, Longwood University

Andrew M. Mountcastle Ph.D. Associate Professor of Biology, Bates College

VII. Approval by the Department Chair (sent via email)

VIII. LITERATURE CITED

1. Alexander, R. McN. (1995). Leg design and jumping technique for humans, other

vertebrates and insects. Philosophical Transactions: Biological Sciences 347(1321):235–

48.

2. Burrows, M. (2009). Jumping performance of planthoppers (Hemiptera, Issidae). Journal

of Experimental Biology 212(17):2844–55.

3. Burrows, M. and Morris, O. (2003). Jumping and kicking in bush crickets. Journal of

Experimental Biology 206(6):1035–49.

4. Burrows, M. and Picker, M. (2010). Jumping mechanisms and performance of pygmy

mole crickets (Orthoptera, Tridactylidae). The Journal of Experimental Biology

213:2386–98.

5. Cross, S. L., Craig, M. D., Tomlinson, S., & Bateman, P. W. (2020). I don’t like crickets,

I love them: Invertebrates are an important Prey Source for Varanid lizards. Journal of

Zoology 310(4):323–33.

6. Hawlena, D., Kress, H., Dufresne, E.R., & Schmitz, O. J. (2011). Grasshoppers alter

jumping biomechanics to enhance escape performance under chronic risk of spider

predation. Functional Ecology 25(1):279–88.

7. Hustert, R. and Baldus, M. (2010). Ballistic movements of jumping legs implemented as

variable components of cricket behaviour. Journal of Experimental Biology

214(12):18112.

8. Li, F., Lin W., Fu, X., Bonsignori, G., Scarfogliero U., Stefanini C., & Dario P. (2012).

Jumping like an insect: design and dynamic optimization of a jumping mini robot based

on bio-mimetic inspiration. Mechatronics 22(2):167–76.

9. Mathis, A., Mamidanna, P., Cury, K. M., Abe, T., Murthy V. N., Mathis, M. W., &

Bethge, M. (2018). DeepLabCut: markerless pose estimation of user-defined body parts

with deep learning. Nature Neuroscience 21(9):1281–89.

10. Patek, S. N., Dudek, D. M., & Rosario, M. V. (2011). From bouncy legs to poisoned

arrows: elastic movements in invertebrates. Journal of Experimental Biology

214(12):1973–80.

11. Sato, N., Shidara, H., & Ogawa, H. (2019). Trade-off between motor performance and

behavioural flexibility in the action selection of cricket escape behaviour. Scientific

Reports 9(1):18112.

12. Tauber, E., and Camhi, J. (1995). The wind-evoked escape behavior of the cricket

Gryllus Bimaculatus : integration of behavioral elements. The Journal of Experimental

Biology 198:1895–1907.

IX. FIGURES

Figure 1. Advantages of having longer hind legs for jumping performance. Long hind legs

result in a longer acting force on substrate, which then results in a higher takeoff velocity and

a greater jump distance produces more work in a jump that aids a prey for a faster predatory

escape.

Figure 2. Basic layout of cricket leg extension. (A) depicts a cricket’s hind leg at resting

point a lower COM. (B) depicts a cricket’s hind leg at full extension a higher COM.

Figure 3. Preliminary tracked data from DeepLabCut of camel cricket jump performance in

lab. Colored dots include: purple (Head), green (Hip), blue (Abrear or lower abdomen),

orange (Knee), and red (Claw). (A) depicting the stance in which the cricket is preparing for a

jump by bring the head and abdomen inwards and pointing hind legs outwards. (B) depicting

the extension of the hind legs at the jump end frame.

Figure 4. Hypothesis of what is predicted that might happen with the percentages of

remaining crickets after predator-prey interaction trials. This is based on the leg lengths of

cave crickets and advantages of higher take off velocity and greater distances to escape

predators.

FIELD CRICKETS CAVE CRICKETS

% REMAINING

HYPOTHESIS OF CRICKETS

REMAINING AFTER PREDATION