Investigating the Mechanism of Organs Related to Sound Production and Reception in Ants

Abstract

Ant colonies require an effective communication system to coordinate the activities of their members. Growing evidence indicates that acoustic communication plays a crucial role in the behavior of ants within a colony. Verse 18 of Surah An-Naml in the Holy Quran also emphasizes verbal interaction among ants in times of danger. This study focuses on investigating the reasons for sound and vibration production and the functioning of sound-producing and receiving organs. Our analysis reveals that sound production in ants occurs through three methods: stridulation, tymbalization, and crepitation. Additionally, characteristics such as duration, frequency spectrum, sound intensity, and environmental conditions of the produced sounds influence the message received by other ants. Furthermore, ants possess the ability to produce and perceive sounds through auditory sensors (antennae), Johnston’s organ, subgenual organs, and mechanical sensors (chordotonal organs, trichoid sensilla structures). Therefore, a research proposal has been developed, encompassing the following stages: 1) Establishing and operating an ant laboratory to create colonies of different ant species under controlled conditions for sound recording and imaging; 2) Conducting studies on native ant species in this field; 3) Imaging sound-producing and receiving organs; 4) Simulating these organs; 5) Recording ant sounds; and finally, 6) Understanding the mechanisms of sound production and reception in ants.

Keywords: Acoustic communication, sound production mechanism, auditory communication.

1. Introduction
The narrative of verbal interaction among ants and its perception by Prophet Solomon (AS), derived from Verse 18 of Surah An-Naml, has been regarded as a legendary belief in Islamic civilizations for centuries. Since ant societies—and insects in general—require an efficient communication system to coordinate their members’ activities, numerous studies have been conducted on their various communication methods. In Verse 18 of Surah An-Naml, Allah the Almighty uses the phrase “Qālat namlatun yā ayyuhan-naml” (An ant said, ‘O ants!’) to illustrate a warning message from one ant to others in the colony. The clarity of this phrase alone is sufficient to guide researchers toward exploring the nature of interaction among ants. The primary objective of this scientific study is to answer the fundamental question of how such communication is possible. Research in this field has identified five communication methods among ants: chemical (pheromones), tactile, body language, trophallaxis (food sharing), and acoustic communication. However, transmitting a message from one ant to a large swarm seems feasible only through pheromones or acoustic means [1]. According to a hadith attributed to Imam Reza (AS) regarding Verse 18 of Surah An-Naml, sound is explicitly mentioned as a communicative factor [2, 3]. Additionally, studies suggest that acoustic interaction is far more plausible for warnings, as it is instantaneously transmitted to all colony members. Based on these explanations, this research initially focuses on sound, acoustic signals, and verbal interaction among these living organisms. As part of a larger proposal, this study seeks to address the following questions: First, how is sound produced and received in ants? Second, what is the phonetic alphabet in ant societies, and how are words defined? Third, what combinations of sentences are constructed using this alphabet, how are conversations conducted, and what types of messages are conveyed? Furthermore, the use of the phrase “Wādi an-naml” (the valley of the ants) by Allah the Almighty is highly significant in this study, as it may influence the selection of ant species and the role of environmental factors in sound transmission within the colony, as well as its reception by Prophet Solomon (AS). As a preliminary step, this study concentrates on the mechanisms of sound production and reception in ants to enable a more effective investigation of the subject.

2. Research Background

Although sound production in ants when threatened or attacked by predators has been reported by various researchers for centuries [4], this subject remains the most contentious issue in studies of this insect species, and its precise mechanism and role in their survival remain unclear [5]. Acoustic communication in ants (like other insects) is divided into two distinct categories depending on the transmission medium:

1. Sound: Acoustic waves transmitted through air.
2. Vibration: Acoustic waves transmitted through substrates (the ground or any surface the ant is on).

Vibrational communication among ants, as with many other insects, is well-established. However, significant disagreements persist among research groups regarding airborne acoustic communication—ranging from those who argue that ants are deaf [6,7] to those who consider acoustic interaction more critical than chemical communication [8].

Research on a species of butterfly larva that mimics the sound of an ant queen has further complicated assumptions about acoustic interactions in ant colonies. This parasitic larva coexists within ant colonies for up to 11 months, attaining a queen-like social status (receiving feeding priority even during food scarcity, preferential relocation during nest migration, and protection priority during predator attacks) [9]. Additionally, a Japanese research team demonstrated that when sound-producing organs or pheromone glands were blocked in farming ants, colonies with disrupted acoustic communication suffered greater agricultural damage (60% loss) compared to those with disrupted pheromonal communication (30% loss). This suggests that sound plays a superior role to chemical signals in the routine communication of fungus-farming ants [10].

Acoustic studies (encompassing both sound and vibration) progress slowly due to data collection and analysis complexities, leaving key questions unresolved. Bioacoustics in insects is an emerging field focusing on sound production, propagation, and reception in insects, with applications in ecological monitoring and species identification. Research on insect acoustic signals employs diverse methods, technologies, and terminologies to decipher these complex messages. Among invertebrates, insects are the only group extensively utilizing airborne sound production and reception. Given these considerations, this study prioritizes:

• Investigating the purposes of sound/vibration production.
• Examining the mechanisms of sound-producing organs.
• Classifying sound-producing/receiving organs.
• Analyzing transmitted sound characteristics.

3. Research Methodology

3-1. Theological and Quranic Research Methodology

After selecting Verse 18 of Surah An-Naml, its meaning was analyzed using authoritative exegeses (Tafsir al-Mizan, Tafsir Nemouneh, Al-Amthal fi Tafsir Kitab Allah al-Munzal, Al-Tibyan fi Tafsir al-Quran, Majma’ al-Bayan, etc.). Key strategic research components were extracted based on the verse’s interpretation, national research priorities, and the team’s expertise. Two foundational concepts were emphasized:

1. “Qālat namlatun yā ayyuhan-naml” (An ant said, “O ants!”): Highlights vocal interaction and warning message transmission.
2. “Wādi an-naml” (The valley of the ants): Indicates geographical and species-specific factors influencing communication.

3-2. Scientific Research Methodology

Due to limited prior work on ant sound-producing organs and acoustic features, this study comprehensively reviews all relevant research from the past 30 years (1995–2024). A systematic search on Science Direct using keywords (Ant, Sound, Communication, Crepitation, Tymbalization, Stridulation, Bioacoustics) identified 62 peer-reviewed articles, of which 38 were selected for their direct relevance to the research objectives.

4. Results, and Discussion

Based on two key phrases extracted from Verse 18 of Surah An-Naml, the research was conducted in two parallel domains:

1. Vocal communication
2. Selection of ant species

For more precise results, the vocal communication domain was divided into four sections:

1. Sound production methods
2. Types of sound-producing organs
3. Function and types of sound-receiving organs
4. Characteristics of transmitted sound

4-1. Sound Production Methods

Among various sound production methods in insects, ants employ three primary mechanisms to generate sound output:

1. Stridulation: Rubbing body parts together as the main method [11]
2. Tymbalization: Striking certain organs against various substrates (observed in Formicinae subfamily) [12,13]
3. Crepitation: Rapid mandible movement and scraping of the lower jaw against substrates [14]

Studies show that the loudest insect sounds are produced through either stridulation or striking organs against substrates or other body parts [4].

Stridulation-produced messages allow ants to transmit information about their status and environmental conditions. Ant interactions through this multimodal communication system, integrating vibrational and chemical messages within their colonies, are highly complex, and the detailed mechanisms remain unclear. The produced sounds are typically emitted during specific social activities including:

• Nest defense [15,16]
• Colony migration [17]
• Nest excavation [18]
• Foraging [19,20]
• Leaf-cutting [14]
• Trophallaxis [21]
• Mating [22]
• Encountering danger and intra/interspecies conflicts [23]

4-2. Types of Sound-Producing Organs

Research has established that ants use stridulatory organs for primary sound production (stridulation), typically located in the abdomen (posterior section of the fourth abdominal segment) [24]. This organ consists of:

• A file-like surface with parallel ridges (pars stridens)
• A scraper (plectrum) in the form of a small protrusion [25,26]

When the ant moves its abdomen, the scraper rubs against the file, producing sound waves often described as “chirping.” For the third method (crepitation), ants use their mandibles to produce various sounds [27,28]. Further studies on different ant species are needed to better understand the sound-producing organs involved in this process.

4-3. Mechanisms and Organs Involved in Sound Reception

The mechanism of sound reception in ants is equally important as sound production. Unlike many insects, ants lack distinct ears. They detect and interpret sound waves through specialized sensors distributed across their body surface and legs. This unique system primarily operates based on receiving vibrational and acoustic signals.

According to studies, ant sensors can be categorized into four groups:

1. Auditory sensors (antennae)
2. Johnston’s organ
3. Subgenual organ
4. Mechanosensors (including trichoid sensilla structures and chordotonal organs)

4-3-1. Auditory Sensors (Antennae)

Antennae serve as primary sensors for detecting airborne vibrations and odors. Using these sensors, ants can detect changes in air pressure and vibrations caused by sounds [29].

4-3-2. Johnston’s Organ

Located within the antennae, Johnston’s organ detects air movements but is insufficiently sensitive for direct perception of stridulatory sounds. However, it plays a role in detecting environmental vibrations [30].

4-3-3. Subgenual Organ

Positioned near the ants’ legs (proximal tibia), this organ primarily detects substrate-borne vibrations and is highly sensitive to low-frequency vibrations [31]. It allows ants to detect subtle environmental changes.

The subgenual organ is particularly important for sensing vibrations transmitted through the ground, while Johnston’s organ responds to air movements and is less effective for stridulatory sounds. These organs are crucial for detecting substrate-borne vibrations.

4-3-4. Mechanosensors

Ant legs and bodies contain mechanosensors that facilitate sound detection, enabling them to sense vibrations from surfaces. These sensors are not adept at detecting airborne sounds over long distances but are more effective in near-field communication where sound characteristics change rapidly over short distances. Mechanosensors allow ants to easily detect substrate-borne sounds and vibrations from other organisms’ movements.

Mechanosensors can be divided into:

1. Chordotonal organs: Function as receivers responsible for detecting sound waves, including both subgenual organs and Johnston’s organ [32]. These organs consist of clusters of sensory units called scolopidia, which vary in number and complexity. They perform both exteroceptive (detecting external stimuli) and proprioceptive (sensing internal body movements) functions. Chordotonal organs are integral to ants’ sensory systems, enabling movement through their environment and effective communication via various mechanical stimuli.
2. Trichoid sensilla: Hair-like structures distributed across the body surface, particularly on antennae, that are rapidly responsive to particle sounds. These allow ants to detect sound waves in the near-field (extending approximately 100 mm around them) [25].

Unlike humans, ants do not respond to airborne sounds over long distances. Instead, they are adapted to sense vibrations and sounds occurring in their immediate vicinity.

4-4. Characteristics of Transmitted Sound

The properties of sounds produced by ants are crucial for understanding their communication systems, as they elucidate the role of airborne sounds alongside substrate-borne vibrations. Comprehending these characteristics is essential for extracting distinct acoustic patterns associated with different roles, conditions, and environmental contexts, ultimately facilitating the interpretation of vocal interactions within ant colonies. Key acoustic parameters include sound intensity (wave amplitude), frequency spectrum, duration, and context-dependent environmental factors, which will be examined in detail below.

4-4-1. Sound Intensity (Wave Amplitude)

Ant acoustic interactions reveal a sophisticated communication system. According to Hicking (2000), ants cannot perceive airborne sounds over long distances (1 meter) and primarily utilize short-range acoustic communication (approximately 10 cm) [25]. The intensity of ant-produced sounds is significantly lower than human auditory thresholds, often requiring amplification for clear detection and recording. The loudest recorded insect communication sounds can reach 110 dB at close range, while quieter signals may measure around 49.8 dB [33]. Sound intensity varies markedly among different ant species and social castes (workers, queens, and males), with queen-produced sound waves exhibiting approximately double the amplitude of males and quadruple that of workers [35].

4-4-2. Frequency Spectrum

Stridulatory sounds produced by ants are typically difficult for humans to detect, yet they are continuously generated at low intensities within colonies. These sounds predominantly fall within the audible frequency range (20 Hz to 20 kHz), with most energy concentrated between 250-4000 Hz. Many species produce sounds near 1000 Hz, though certain ants demonstrate broader spectral characteristics. Massoni et al. (2021) evaluated frequency spectra in leafcutter ants (Atta cephalotes), revealing size-dependent variations: smaller workers produced stridulatory frequencies between 2000-5000 Hz, whereas major workers generated ultra-high frequencies of 38,000-46,000 Hz [35]. This demonstrates that spectral characteristics are influenced by both body size and stridulatory organ morphology across species [36]. Furthermore, distinct frequency profiles were associated with different behavioral states – alarm signals exhibited the highest frequencies, while awareness signals occupied the lowest spectral range. Food source quality and environmental conditions were also found to modulate spectral properties.

4-4-3. Duration

During stridulation, insects perform periodic mechanical movements that generate oscillatory vibrations. These vibrations typically exhibit a cyclical pattern, producing a fundamental frequency along with harmonic components (integer multiples of the fundamental frequency) that may be selectively amplified or attenuated depending on the insect’s anatomical structure and sound production mechanism [37].

Temporal patterns of sound emission vary according to behavioral context (e.g., foraging vs. alarm signals) and species-specific characteristics. In black ants, stridulatory signals persist for differing durations depending on their communicative function: alarm signals remain detectable for 10 minutes, alert/attack signals for 1-3 minutes, and distress signals for 4 minutes before dissipating [25]. Individual signal duration averages 8 seconds across contexts [25].

4-4-4. Context-Dependent Environmental Factors

Atmospheric conditions (temperature, humidity, air pressure) and elevation above ground level influence airborne sound propagation. Additionally, ants generate substrate-borne vibrations through whole-body movements or specific appendage strikes, which interact variably with different substrates (soil, wood, leaves, water, artificial materials). Denser materials generally support more efficient vibrational transmission, though studies demonstrate effective propagation of mandible-generated mechanical sounds across porous or soft substrates [38]. This substrate dependence plays a critical role in ant vibrational communication systems.

4-5. Second Domain: Target Ant Species Selection

The selection of appropriate ant species constitutes another critical research consideration. While most existing studies focus on leafcutter ants (Atta spp.) and other American species, their limited accessibility and the study’s primary objective – investigating the Quranic account in Surah An-Naml (27:18) – necessitate alternative approaches.

The Quranic phrase “وَادِ النَّمْلِ” (Valley of the Ants) provides key selection criteria, suggesting:

1. Large-nesting species: Colonies substantial enough to constitute identifiable “valleys”
2. Geographical context: Likely species inhabiting historical Saba (Sheba) region

Cross-referencing subsequent verses (27:20-24) regarding Prophet Solomon’s hoopoe (hudhud) provides additional geographical markers. Historical and exegetical sources [39-41] locate ancient Saba in modern Yemen, with its capital Marib approximately 120 km east of Sana’a. By estimating the hoopoe’s daily flight range from Marib and focusing on large-colony species within this radius, potential target species can be identified.

“Ants employ three primary sound production mechanisms (stridulation, tymbalization, crepitation) detected through specialized sensory organs. This sophisticated communication system plays a vital role in collective behavior coordination and may inspire artificial communication systems.”

5. Conclusions and Directions for Future Research

Based on comprehensive analyses of recent scientific studies and the Quranic verse describing the detailed, sequential transmission of warning signals (regarding the arrival of Prophet Solomon and his army) from one individual to an entire community, extensive research on sound production and reception in ants is warranted. This research demands a multidimensional approach and should be conducted in collaboration with the research groups proposed in the results section. Accordingly, a research proposal has been developed with the following stages:

1. Establishment of an Ant Laboratory:
   1. Priority should be given to setting up a specialized ant laboratory.
   1. Constructing controlled colonies of various ant species for sound recording and imaging is essential.
   1. Acquiring specialized bioacoustic equipment for measuring both airborne and substrate-borne vibrations at very low intensities across different frequency spectra is necessary.
2. Replication of Existing Studies on Native Species:
   1. Initial efforts will focus on replicating existing studies on sound production in native Iranian ant species, followed by species from regions near Yemen.
   1. Studies on ant species from these two biomes will be prioritized, with at least three dominant species selected from each.
   1. Iran’s biome is preferred due to easier access and sample preparation, while areas near Yemen are significant due to the narrative in Verses 18–22 of Surah An-Naml.
   1. An appropriate number of individuals from each species will be included for both individual and population-level studies.
3. High-Resolution Imaging of Sound-Related Organs:
   1. High-quality imaging of sound-producing and receiving organs will be performed using electron microscopy on specimens prepared under standardized protocols.
   1. Intra- and interspecies comparisons of these organs will be thoroughly documented.
4. 3D Modeling of Sound-Producing and Sensory Organs:
   1. Based on the obtained images, we aim to develop three-dimensional models of sound-producing organs and sensory structures.
   1. These models will enable simulations of sound production, propagation, and reception, leading to a deeper understanding of bioacoustic behaviors in ants.
5. Sound Recording Under Varied Conditions:
   1. Recording sounds in different environmental contexts is a critical component of this research.
   1. Investigations will include sounds produced by individuals from different social castes, isolated vocalizations, and colony-wide acoustic signals.
6. Anatomical-Functional Correlation of Sound-Related Organs:
   1. The final phase will explore the relationship between the anatomy of sound-producing/receiving organs and their functional mechanisms.
   1. This is particularly crucial for ant auditory sensors, which remain understudied.

Proposed Collaborative Research Groups

Research on sound production and reception in ants (both substrate-borne and airborne) has been conducted by various teams worldwide. Four prominent research groups have published groundbreaking studies and are recommended for future collaborations:

1. Barbero Group (Italy):
   1. Extensive research on ant bioacoustics, with a primary focus on substrate-borne sound production and reception.
2. Japanese Research Team (Murakami, Sakamoto, and Higashi):
   1. Investigated the selective blocking of acoustic and pheromonal communication, demonstrating the relative importance of each modality in maintaining colony function.
3. Russian Research Group (Led by Prof. Zhanna Reznikova):
   1. Published seminal works on vocal communication among ants [42–46].
4. Hickling Research Group:
   1. Focused on acoustic signal characteristics, including intensity, frequency spectrum, temporal patterns, and context-dependent features.
   1. Recommended as a key collaborator for future research initiatives.

This structured approach will advance our understanding of ant communication systems while bridging scientific inquiry with Quranic insights. Collaborative efforts with leading research teams will ensure methodological rigor and interdisciplinary innovation.

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