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Swallowing

Swallowing, also known as deglutition, is the complex physiological process that transports ingested substances from the mouth (oral cavity) to the stomach via the pharynx and esophagus, while protecting the airway from aspiration and facilitating digestion.^[1] This essential function involves the coordinated activity of over 30 muscles and multiple cranial nerves, combining voluntary and reflexive actions to form a bolus of food or liquid and propel it safely through the upper digestive tract.^[2] Biologically, swallowing is crucial for nutrition, hydration, and preventing life-threatening complications like aspiration pneumonia, underpinning survival across species. Evolutionarily, it has developed over millions of years in vertebrates; the modern human and mammalian form features a permanent intersection of respiratory and digestive pathways, enabling precise control and reduced aspiration risk compared to earlier suction-based mechanisms in fish and amphibians.^[3] Swallowing begins as early as 15 weeks in utero and is refined through infancy, with adults performing it approximately 600–1,000 times per day, often subconsciously.^[1] The swallowing process is typically divided into three main phases: oral, pharyngeal, and esophageal, each with distinct anatomical and physiological mechanisms.^[1] The oral phase is voluntary and preparatory, where the tongue manipulates and propels the bolus toward the oropharynx at speeds of about 4–20 cm/s, involving muscles such as the genioglossus and hyoglossus innervated primarily by the hypoglossal nerve (cranial nerve XII).^[1] This phase seals the bolus to prevent premature spillage and includes mastication for solids, adapting to bolus consistency—liquids require less preparation than solids.^[2] Once initiated, the pharyngeal phase becomes reflexive and irreversible, lasting about 1 second and transporting the bolus through the pharynx at 20–40 cm/s while safeguarding the airway.^[1] Key actions include elevation of the hyoid bone and larynx, closure of the nasopharynx via the soft palate, and airway protection through epiglottis inversion, vocal fold adduction, and transient apnea (0.5–1.5 seconds), coordinated by cranial nerves IX, X, and the nucleus ambiguus in the brainstem.^[2] The upper esophageal sphincter (UES), composed mainly of the cricopharyngeus muscle, relaxes to allow passage, opening via hyoid elevation and bolus pressure.^[1] The esophageal phase is primarily involuntary, relying on sequential peristaltic contractions to move the bolus to the stomach at 3–4 cm/s, aided by gravity in upright positions.^[1] Primary peristalsis is triggered by the swallow, while secondary peristalsis clears residue, involving smooth muscle from the upper third transitioning to the lower two-thirds, with the lower esophageal sphincter (LES) relaxing to permit entry into the stomach.^[2] Disruptions in any phase can lead to dysphagia, highlighting swallowing's role in nutrition, hydration, and overall health.^[1]

Overview and Importance

Definition and Process Summary

Swallowing, also known as deglutition, is the physiological process of transporting ingested substances such as food, liquids, or saliva from the oral cavity to the stomach via the pharynx and esophagus through coordinated muscle contractions and relaxations.^[1] This intricate mechanism ensures the safe and efficient movement of boluses while protecting vital structures like the airway.^[4] The swallowing process unfolds in three primary phases: the oral phase, which involves voluntary manipulation and propulsion of the bolus; the pharyngeal phase, an involuntary reflex that propels the bolus into the esophagus; and the esophageal phase, which facilitates peristaltic transport to the stomach.^[1] These phases occur as a seamless, sequential event, typically lasting less than 10 seconds in healthy adults, with the initial voluntary trigger giving way to reflexive coordination.^[2] Swallowing is crucial for maintaining nutrition and hydration by enabling the ingestion of essential calories and fluids, while also initiating digestion through gastric delivery of substrates.^[1] It critically prevents aspiration by sealing the airway during bolus passage, thereby averting respiratory complications such as pneumonia.^[2] The understanding of swallowing impairments has ancient roots, with the Hippocratic school recognizing dysphagia in patients suffering head trauma, often associating it with neurological deficits and secondary infections.^[5]

Biological and Evolutionary Role

Swallowing represents an ancient physiological process integral to the evolution of vertebrate digestive systems, originating over 570 million years ago in the common ancestor of chordates and hemichordates with the development of pharyngeal structures from gill-like apparatuses. Genomic evidence from acorn worm genomes illustrates how these primitive pharyngeal slits evolved into the foundational aerodigestive tract, enabling the transition from filter-feeding in invertebrates to more active ingestion in early vertebrates.^[6] By approximately 350 million years ago, the emergence of a muscular tongue in sarcopterygian fish marked a pivotal shift from suction-based feeding—common in aquatic species—to tongue-propelled bolus formation, facilitating the exploitation of terrestrial food sources as vertebrates transitioned to tetrapods.^[7] This evolutionary progression enhanced feeding efficiency by allowing precise food manipulation and transport, underscoring swallowing's role as a conserved trait for survival across diverse environments. In biological terms, swallowing primarily facilitates nutrient intake by propelling food boluses through the digestive tract while simultaneously protecting the airway through coordinated closure of the glottis and epiglottis, preventing aspiration and ensuring respiratory safety during feeding.^[2] These dual functions—ingestion and protection—have been conserved since early vertebrate evolution, adapting to varying ecological pressures. In mammals, a unique duality emerges where the same hyoid-tongue apparatus supports both swallowing and vocalization; this shared neural and muscular framework, evident in suckling and mastication systems, originated around 165 million years ago in Jurassic mammals like Microdocodon gracilis, whose flexible hyoid bones enabled sophisticated bolus propulsion akin to modern forms.^[8]^[9] This mammalian innovation not only optimized feeding but also laid the groundwork for advanced communication in humans, where swallowing and speech compete for the same oropharyngeal space.^[10] Comparative analyses reveal variations in swallowing efficiency linked to dietary adaptations, with carnivores exhibiting faster bolus transit to accommodate rapid ingestion of whole prey, as seen in their streamlined pharyngeal propulsion compared to the more deliberate, multi-bolus processes in herbivores processing fibrous plant matter.^[11] For instance, mammalian carnivores achieve quicker deglutition cycles to minimize exposure during predation, enhancing survival rates, whereas herbivores invest in extended oral processing for optimal nutrient extraction from recalcitrant diets.^[12] These differences highlight swallowing's adaptive plasticity, where speed and bolus size correlate with ecological niches, from the swift swallows of piscivorous fish to the efficient, volume-handling mechanisms in grazing mammals. In contemporary human contexts, effective swallowing plays a critical role in preventing malnutrition by ensuring adequate caloric and nutrient absorption, particularly in vulnerable populations where impairments can lead to reduced intake and weight loss.^[13] Aging-related declines in swallowing function, such as weakened pharyngeal muscles and delayed reflexes starting around age 60, contribute to frailty and nutritional deficits, exacerbating risks of sarcopenia and overall health deterioration.^[14] Interventions preserving swallowing integrity, like targeted rehabilitation, thus mitigate these declines, supporting longevity and quality of life.^[15]

Anatomy of Swallowing

Key Structures in the Oral Cavity

The oral cavity serves as the initial site for the preparatory and transit phases of swallowing, where food is manipulated into a cohesive bolus suitable for propulsion toward the pharynx. This region is bounded anteriorly and laterally by the lips and cheeks, superiorly by the hard and soft palates, inferiorly by the mylohyoid muscles forming the floor of the mouth, and posteriorly by the fauces. Key structures within the oral cavity facilitate mastication, bolus formation, lubrication, and controlled posterior movement, all coordinated by sensory feedback from the trigeminal (CN V), glossopharyngeal (CN IX), and vagus (CN X) nerves.^[16]^[4] The lips, composed of orbicularis oris muscle (innervated by CN VII), form an anterior seal to contain the bolus during the oral preparatory phase, preventing premature spillage and aiding in initial food intake.^[1]^[2] In the oral transit phase, they maintain closure to support tongue-driven propulsion. The teeth, embedded in the alveolar processes of the maxilla and mandible, grind and fragment solid foods during mastication, with assistance from masticatory muscles such as the masseter, temporalis, and medial/lateral pterygoids (all innervated by CN V3). This mechanical breakdown is essential for creating a cohesive bolus of swallowable size, typically 5–25 mL depending on consistency.^[4]^[16]^[17] The tongue is the primary effector organ in the oral cavity for swallowing, consisting of intrinsic and extrinsic muscles that enable precise manipulation. Extrinsic muscles like the genioglossus (protrusion, CN XII), hyoglossus (depression and retraction, CN XII), styloglossus (retraction and elevation, CN XII), and palatoglossus (elevation, CN X) coordinate to gather, compress, and propel the bolus posteriorly against the hard palate during the oral propulsive phase.^[4]^[1] The tongue's dorsum contacts the hard palate to form a posterior seal, ensuring unidirectional bolus flow while sensory receptors detect bolus readiness.^[2] The hard palate, formed by the palatine processes of the maxilla and horizontal plates of the palatine bones, provides a firm, immobile surface against which the tongue squeezes the bolus for posterior transport. Posteriorly, the soft palate (velum), including the uvula, elevates via the tensor veli palatini (CN V3) and levator veli palatini (CN IX/X via pharyngeal plexus) to seal the nasopharynx, preventing nasal regurgitation during swallowing initiation.^[4]^[1] The cheeks, supported by buccinator muscle (CN VII), laterally contain the bolus and assist in positioning food onto the occlusal surfaces of the teeth during mastication.^[2] The floor of the mouth, elevated by the mylohyoid (CN V3) and geniohyoid (C1 via hypoglossal nerve) muscles, supports tongue elevation and contributes to bolus containment.^[1] Salivary glands—parotid (serous, CN IX), submandibular (mixed, CN VII), and sublingual (mucoid, CN VII)—secrete saliva at rates of 0.5-1.5 L/day to moisten and lubricate the bolus, initiating starch digestion and facilitating smooth transit through the oral cavity. This lubrication is critical for reducing friction and ensuring bolus cohesion before pharyngeal transfer.^[16]^[4]

Pharyngeal and Esophageal Structures

The pharynx is a muscular tube extending from the base of the skull to the level of the cricoid cartilage, divided into three regions critical for the pharyngeal phase of swallowing: the nasopharynx (posterior to the nasal cavity, above the soft palate), oropharynx (posterior to the oral cavity, extending to the level of the hyoid bone), and laryngopharynx (also known as hypopharynx, from the hyoid to the esophagus, encompassing the pyriform sinuses and valleculae for bolus containment).^[2] These divisions facilitate the sequential propulsion and protection during bolus transit, with the nasopharynx sealed by palatal elevation to prevent nasal reflux.^[4] The epiglottis, a leaf-shaped elastic cartilage arising from the thyroid cartilage and attached to the hyoid bone via the thyroepiglottic ligament, plays a pivotal role in airway protection by inverting over the laryngeal inlet during swallowing to direct the bolus toward the esophagus.^[2] Pharyngeal muscles include the constrictor group—superior (originating from the pterygomandibular raphe and medial pterygoid plate, inserting into the median raphe), middle (from the hyoid cornua to the median raphe), and inferior (from the cricoid and thyroid cartilages to the esophagus)—which contract sequentially in a peristaltic wave to propel the bolus inferiorly.^[4] Elevators such as the stylopharyngeus (innervated by cranial nerve IX, elevating the pharynx and larynx) and suprahyoid muscles (e.g., mylohyoid, geniohyoid) contribute to hyolaryngeal excursion, lifting the hyoid bone and larynx anteriorly and superiorly to facilitate upper esophageal sphincter opening.^[2] The esophagus is a 25-cm muscular tube connecting the pharynx to the stomach, beginning at the cricoid cartilage (C6 level) and ending at the gastroesophageal junction (T11 level).^[18] Its upper esophageal sphincter (UES), formed primarily by the cricopharyngeus muscle (a component of the inferior pharyngeal constrictor), maintains tonic closure at rest (pressure ~30-100 mmHg) and relaxes via inhibition during swallowing to allow bolus entry.^[2] The esophageal body transitions from striated muscle in the upper third (under somatic control) to smooth muscle in the lower two-thirds (under autonomic control), enabling primary and secondary peristalsis for bolus transport at speeds of 2-4 cm/s.^[18] The lower esophageal sphincter (LES), a high-pressure zone (~15-30 mmHg) at the distal end without a distinct anatomical muscle, relaxes tonically during swallowing to permit gastric entry while preventing reflux.^[1] Supporting structures include the hyoid bone, a U-shaped mobile bone suspended by stylohyoid and digastric muscles and connected to the larynx via the thyrohyoid membrane, which elevates ~2-3 cm during swallowing to tension the pharynx and open the UES.^[2] Laryngeal suspension via suprahyoid and infrahyoid muscles ensures coordinated movement for airway safeguarding. Mucosal folds (longitudinal in the esophagus) and submucosal glands (providing seromucous lubrication) reduce friction and aid bolus passage, with the pharyngeal mucosa featuring stratified squamous epithelium for resilience.^[18] Vascular supply to the pharynx derives from branches of the external carotid (ascending pharyngeal artery) and subclavian (inferior thyroid artery) arteries, forming an anastomotic network along the posterior wall, while the esophagus receives inferior thyroid arteries superiorly and esophageal branches of the aorta inferiorly.^[19] Innervation involves the pharyngeal plexus (from cranial nerves IX and X) for sensory and motor functions, with the esophagus primarily vagal (CN X) via recurrent laryngeal branches; the recurrent laryngeal nerve, looping under the subclavian artery (right) or aortic arch (left), runs in close proximity to the carotid sheath, rendering it vulnerable to injury during neck procedures and potentially causing dysphagia or aspiration.^[4]^[19]

Physiology in Humans

Neural Control and Reflexes

Swallowing is regulated by a central pattern generator (CPG) located in the medulla oblongata, which coordinates the rhythmic motor patterns essential for the process.^[20] This CPG primarily involves the nucleus tractus solitarius (NTS), responsible for sensory integration and pattern generation, and the nucleus ambiguus, which provides motor output to pharyngeal and laryngeal muscles.^[21] The swallowing center is bilaterally represented in the dorsal medulla, ensuring robust neural coordination even if one side is compromised.^[22] Voluntary initiation of swallowing originates from cortical areas, particularly the primary motor and sensory cortices, which send inputs via the corticobulbar tracts to the brainstem swallowing center.^[23] These tracts allow for conscious control, especially during the oral phase, where bolus formation and propulsion are under deliberate regulation.^[24] Once triggered, the transition to reflexive phases occurs rapidly, with the CPG taking over to orchestrate involuntary sequences without further cortical involvement.^[25] Reflex arcs form the core of swallowing's sensory-motor feedback loop, with afferent signals from mechanoreceptors and chemoreceptors in the oropharynx and larynx traveling primarily via the glossopharyngeal (IX), vagus (X), and trigeminal (V) nerves to the NTS.^[26] These inputs detect bolus presence and trigger the reflex, while efferent motor commands are distributed through the vagus nerve (X) for pharyngeal and esophageal muscles and the hypoglossal nerve (XII) for tongue movements.^[22] This circuitry ensures precise timing, with the pharyngeal phase lasting approximately 1 second, during which transient apnea (0.5 to 1.5 seconds) protects the airway during bolus transit.^[1] The oral phase relies on voluntary neural drive for manipulation and transport, contrasting with the involuntary pharyngeal and esophageal phases, which are driven by brainstem reflexes to prevent aspiration and facilitate peristalsis.^[27] Swallowing transiently inhibits respiration at the brainstem level, inducing a brief apnea that aligns with the pharyngeal phase to safeguard the airway, typically interrupting expiration.^[28] Aging impacts neural control by reducing efficiency in central processing and sensory feedback, leading to delayed pharyngeal swallow onset and diminished activation in swallowing-related cortical and brainstem networks.^[29] These changes, including weaker corticobulbar connections and slower reflex arcs, increase vulnerability to inefficient coordination without necessarily causing overt dysfunction in healthy individuals.^[30]

Oral Phase Mechanics

The oral phase of swallowing begins with the voluntary process of bolus preparation, where ingested food or liquid is transformed into a cohesive mass suitable for safe transit. Mastication reduces the particle size of solid foods through rhythmic chewing movements coordinated by the tongue, cheeks, and jaw, softening the material and breaking it down mechanically.^[2] Simultaneously, saliva is mixed with the food particles via glandular secretions from the submandibular, sublingual, and parotid glands, which lubricate and bind the components into a cohesive bolus while initiating enzymatic digestion.^[2] The tongue plays a central role in shaping the bolus by pressing it against the hard palate, manipulating it laterally and posteriorly to ensure uniformity and containment within the oral cavity, preventing premature spillage.^[31] Propulsion during the oral phase involves a sequential biomechanical sequence driven by the tongue's voluntary movements. The tongue tip elevates to contact the hard palate, followed by a wave-like posterior expansion of this contact, which sequentially squeezes the bolus backward along the palatal surface toward the oropharynx.^[32] This posterior thrust propels the bolus through the fauces into the posterior oral cavity, with peak efficiency occurring when the upper and lower teeth are in close approximation.^[2] Concurrently, the soft palate contacts the tongue to seal the oral cavity posteriorly, preventing premature spillage into the oropharynx.^[2] The duration of the oral propulsive phase is typically around 1 second in healthy adults, though it exhibits significant variability based on bolus characteristics. For liquids, transit times range from 0.35 to 1.54 seconds, while pasty foods take 0.39 to 1.05 seconds, and solids can extend from 1 to 12.8 seconds due to extended mastication needs.^[33] This variability reflects adaptations to food texture, with denser or larger boluses requiring prolonged tongue manipulation for adequate preparation and propulsion.^[33] Sensory feedback integrates throughout the oral phase to monitor bolus readiness and position, ensuring coordinated transition to the pharyngeal stage. Mechanoreceptors in the oral mucosa and tongue detect bolus size, consistency, and posterior positioning, providing afferent input via the trigeminal and glossopharyngeal nerves to central pattern generators.^[26] Once the bolus accumulates sufficiently in the oropharynx—typically signaling completion of oral transit—this sensory detection triggers the involuntary pharyngeal swallow reflex, modulating timing based on bolus properties like volume and texture.^[26]

Pharyngeal Phase Dynamics

The pharyngeal phase of swallowing is a rapid, involuntary process triggered by sensory input from the bolus arriving in the oropharynx, initiating a coordinated sequence to propel the bolus toward the esophagus while safeguarding the airway. This phase begins with laryngeal elevation, driven by contraction of the suprahyoid muscles, which pulls the hyoid bone and larynx upward and forward, simultaneously shortening and widening the pharynx to facilitate bolus passage. Concurrently, the epiglottis inverts or retroflexes through passive movement influenced by hyolaryngeal excursion and tongue base retraction, directing the bolus laterally into the piriform sinuses and away from the laryngeal inlet. Vocal fold adduction follows, mediated by the lateral cricoarytenoid and interarytenoid muscles, which approximate the true vocal cords to seal the glottis internally. Finally, a pharyngeal constriction wave propagates via sequential contraction of the superior, middle, and inferior pharyngeal constrictor muscles, generating a peristaltic force at speeds of 20-40 cm/s to strip the bolus residue from the pharyngeal walls and drive it inferiorly.^[1]^[34]^[35] Airway protection during this phase is multifaceted and critical, given the anatomical overlap between the digestive and respiratory tracts. Transient apnea occurs as respiration is inhibited for approximately 0.5-1.5 seconds, primarily during the expiratory phase, to prevent bolus entry into the trachea; this reflex is mediated by central pattern generators in the brainstem. The upper esophageal sphincter (UES) relaxes synchronously, facilitated by traction from hyolaryngeal elevation, inhibition of the cricopharyngeus muscle, and hydrostatic pressure from the incoming bolus, opening the sphincter to a diameter of about 1.5-2 cm for unimpeded transit. These mechanisms ensure minimal residue, but any discoordination—such as delayed elevation or incomplete adduction—heightens the risk of misdirection, potentially leading to penetration, aspiration, or pharyngeal residue, which underscores the phase's irreversible nature once initiated.^[1]^[36]^[34] The entire pharyngeal phase typically lasts 0.5-1 second in healthy adults, reflecting its high-speed execution under brainstem control to minimize exposure time for airway compromise. Differences in bolus consistency influence dynamics within this brief window: liquids, being less viscous, advance more rapidly into the pharynx via momentum and gravity, often reaching the valleculae or hypopharynx before full swallow initiation, which demands precise timing to avoid premature spillage and aspiration. In contrast, solids form a more cohesive bolus requiring greater reliance on the pharyngeal constriction wave for propulsion, as their higher viscosity resists passive flow and necessitates stronger muscular stripping to clear residue.^[1]^[37]^[2]

Esophageal Phase Function

The esophageal phase of swallowing commences immediately following the pharyngeal phase, with the bolus entering the esophagus via relaxation of the upper esophageal sphincter (UES). This phase is involuntary and primarily driven by peristaltic contractions that propel the bolus distally toward the stomach. Primary peristalsis, triggered by the swallowing center in the brainstem, initiates a coordinated wave of muscle contraction along the esophageal body, ensuring efficient transport of the bolus.^[38]^[39] The esophagus is divided into distinct muscular regions: the proximal one-third comprises striated muscle under somatic nervous control via the vagus nerve, facilitating rapid initial propulsion, while the distal two-thirds consists of smooth muscle regulated by the autonomic nervous system through the myenteric plexus, enabling sustained peristaltic activity. Secondary peristalsis, elicited by local distension of the esophageal wall rather than central input, serves as a clearance mechanism to eliminate any residual bolus material that primary peristalsis may leave behind, preventing stagnation. The UES, formed mainly by the cricopharyngeus muscle, relaxes actively during swallowing to permit bolus entry and then contracts to guard against reflux from the esophagus; similarly, the lower esophageal sphincter (LES), a high-pressure zone of smooth muscle augmented by the crural diaphragm, relaxes transiently via inhibitory neurotransmitters like nitric oxide to allow passage into the stomach while maintaining tonic closure to retain gastric contents.^[38]^[2]^[39] This phase typically lasts 8 to 10 seconds for solid boluses, with peristaltic velocity of 3 to 4 cm/s along the approximately 25 cm esophageal length, though liquids transit more rapidly, often in 5 to 6 seconds, due to lower viscosity. Clearance of residue is further supported by secondary peristaltic waves, which can be repeated as needed to ensure complete emptying. Influences such as gravity and posture modulate transport efficiency: in an upright position, gravity assists bolus descent, particularly for liquids, while supine postures increase reliance on peristalsis; intrathoracic pressure variations during respiration and swallowing also affect esophageal dynamics by altering the pressure gradient.^[39]^[2]^[38]

Clinical Aspects

Swallowing Disorders (Dysphagia)

Dysphagia refers to impairments in the swallowing process that disrupt the safe and efficient transport of food, liquids, or saliva from the mouth to the stomach. It is broadly classified into oropharyngeal dysphagia, which involves difficulties in the oral and pharyngeal phases often linked to neurological or muscular dysfunction, and esophageal dysphagia, which affects the esophageal phase due to motility or structural abnormalities. Oropharyngeal dysphagia commonly arises from neurogenic causes such as stroke, where up to 51-73% of patients experience swallowing impairments due to disrupted neural coordination.^[40] Esophageal dysphagia, in contrast, frequently results from structural issues like strictures or rings that narrow the esophageal lumen, impeding bolus passage.^[40] Overall etiologies encompass neurogenic factors (e.g., central or peripheral nervous system disorders), structural anomalies (e.g., tumors or inflammation), and iatrogenic origins (e.g., complications from surgeries or medications).^[41] Common swallowing disorders include achalasia, characterized by failed esophageal peristalsis and incomplete relaxation of the lower esophageal sphincter, leading to progressive dysphagia for both solids and liquids.^[42] Globus sensation presents as a persistent, non-painful feeling of a lump in the throat without actual obstruction or true swallowing difficulty, often associated with heightened pharyngeal sensitivity rather than mechanical issues.^[43] A significant complication across disorders is the risk of aspiration pneumonia, particularly from silent aspiration where material enters the airway without overt coughing or choking; this occurs in about half of dysphagia cases and elevates pneumonia risk threefold.^[44]^[45] Symptoms of dysphagia vary by type but commonly include coughing or choking during meals, a sensation of food sticking in the throat or chest, painful swallowing (odynophagia), nasal regurgitation, and unintended weight loss due to reduced intake.^[40] In oropharyngeal cases, additional signs like drooling or a wet voice may occur from delayed bolus clearance. Prevalence is notably high among the elderly, affecting 40-60% of nursing home residents, with pooled estimates reaching 56% based on screening tools.^[46]^[47] Key risk factors include neurological conditions such as Parkinson's disease, where basal ganglia lesions impair swallow initiation in up to 80% of advanced cases, and amyotrophic lateral sclerosis (ALS), which causes bulbar muscle weakness leading to dysphagia in most patients by disease progression.^[41]^[48] Post-surgical complications, such as those following head and neck procedures or tracheostomy (with 50-83% aspiration risk), further exacerbate vulnerability.^[40] Age-related factors like sarcopenia-induced muscle weakness and reduced salivary flow also heighten susceptibility, particularly in those over 65.^[49]

Diagnosis and Management

Diagnosis of swallowing disorders, or dysphagia, typically involves a combination of clinical assessments and instrumental evaluations to identify impairments in the oral, pharyngeal, or esophageal phases. The videofluoroscopic swallow study (VFSS), also known as a modified barium swallow, is a radiographic procedure that visualizes the swallowing process in real-time using fluoroscopy, allowing clinicians to detect abnormalities such as aspiration or residue.^[50] VFSS is considered the gold standard for evaluating oropharyngeal dysphagia across all ages, as it distinguishes anatomic and physiologic causes of impairment.^[51] Another key technique is the fiberoptic endoscopic evaluation of swallowing (FEES), a portable endoscopic procedure performed transnasally at the bedside or in clinic, which directly observes laryngeal and pharyngeal structures during swallowing to assess secretion management, sensation, and bolus flow.^[52] FEES is particularly useful for patients unable to undergo radiation exposure or transport, providing immediate feedback on safe oral intake levels.^[53] For esophageal disorders, high-resolution manometry measures intraluminal pressures and motility patterns during swallows, aiding diagnosis of conditions like achalasia by evaluating sphincter relaxation and peristalsis.^[54] This test is essential for planning interventions in non-obstructive dysphagia.^[55] Management of dysphagia emphasizes a multidisciplinary approach involving speech-language pathologists (SLPs), gastroenterologists, otolaryngologists, and nutritionists to tailor interventions based on diagnostic findings. SLPs lead rehabilitative swallowing therapy, such as the Mendelsohn maneuver, which involves voluntary prolongation of laryngeal elevation during the swallow to enhance hyolaryngeal excursion, upper esophageal sphincter opening, and bolus clearance.^[56] This exercise has been shown to improve pharyngeal pressures and reduce residue in patients with reduced laryngeal movement.^[57] Dietary modifications, including the use of commercial thickeners to increase liquid viscosity, help control bolus flow and minimize aspiration risk, particularly in neurogenic dysphagia.^[58] For severe esophageal motility issues like achalasia, surgical options such as laparoscopic Heller myotomy cut the lower esophageal sphincter muscles to facilitate passage of food, often combined with an antireflux procedure for long-term symptom relief.^[59] Peroral endoscopic myotomy (POEM) represents a less invasive alternative, performed endoscopically with comparable efficacy.^[60] Effective management yields measurable outcomes, including reduced aspiration rates and improved quality of life. Swallowing therapy interventions, such as exercises combined with postural techniques, have demonstrated decreased penetration-aspiration scores and shorter hospital stays in stroke patients.^[61] Thickened liquids, for instance, lower aspiration incidence to as low as 8.3% compared to thin liquids in hospitalized individuals with dysphagia.^[62] Multidisciplinary protocols integrating therapy and modifications can reduce chest infections and enhance oral intake, with aspiration prevention surgeries further decreasing suctioning needs and caregiver burden.^[63] As of 2025, recent advances include AI-assisted imaging for early detection, where deep learning algorithms analyze VFSS videos to automatically quantify dysphagia severity, improving diagnostic accuracy and enabling personalized rehabilitation.^[64] Neuromodulation devices, such as repetitive transcranial magnetic stimulation (rTMS) and vagal nerve magnetic stimulation, target central and peripheral pathways to enhance swallowing function in post-stroke and neurogenic cases, showing promise in reducing cricopharyngeal dysfunction through non-invasive neural plasticity.^[65] These technologies facilitate faster, more precise interventions within multidisciplinary frameworks.^[66]

Comparative Swallowing

In Other Mammals

Swallowing in non-human mammals exhibits significant adaptations tied to dietary habits, body size, and ecological niches, reflecting evolutionary pressures for efficient nutrient acquisition and survival. In ruminants like cows, the process involves initial rapid swallowing of forage into the rumen for microbial fermentation, followed by regurgitation, remastication, and reswallowing during rumination to break down fibrous material. This cyclic mechanism enhances digestibility of plant-based diets, with regurgitation triggered by reticular contractions and esophageal sphincter relaxation to propel boluses back to the mouth.^[67]^[68] In contrast, carnivores demonstrate streamlined swallowing suited to whole-prey consumption, where large boluses of meat are transported quickly through the pharynx and esophagus with minimal mastication. Anatomical variations further underscore these dietary specializations. Herbivores often possess longer esophagi relative to body size to accommodate transport to specialized foregut chambers; for instance, in donkeys, the esophagus measures 89–110 cm, facilitating efficient delivery of bulky plant matter to fermentation sites.^[69] Primates, including non-human species like macaques, feature enhanced tongue mobility due to a more flexible hyolingual apparatus, enabling precise manipulation and propulsion of diverse food items during the oral phase of swallowing, which supports omnivorous diets and tool use.^[70] Physiological adjustments also scale with size: small mammals, such as rats, exhibit accelerated pharyngeal phases, compared to longer durations in larger species, allowing rapid ingestion to evade predators.^[71] Neonatal mammals across taxa rely on coordinated suckle-swallow-breathe patterns for milk intake, where rhythmic sucking generates negative pressure to draw fluid, followed by pharyngeal swallowing and expiratory pauses to prevent aspiration, a pattern present from birth in species like pigs and extending into early development.^[72] These adaptations highlight how mammalian swallowing balances ingestion efficiency with respiratory and sensory demands.

In Non-Mammalian Animals

In non-mammalian vertebrates, swallowing mechanisms vary widely to accommodate diverse aquatic and terrestrial environments, often integrating feeding with respiration. Fish, for instance, employ buccal pumping, where rhythmic expansion and contraction of the buccal cavity draw water and food into the mouth, followed by pharyngeal jaw manipulation to transport the bolus toward the esophagus.^[73] Unlike mammals, fish lack a true epiglottis, relying instead on opercular movements and branchial arches to direct food while ventilating gills, with peristalsis initiating in the esophagus without a distinct pharyngeal phase.^[73] This system is evident in species like channel catfish, where X-ray reconstruction of moving morphology reveals coordinated hyoid and jaw motions propelling food posteriorly in discrete swallows.^[73] Birds exhibit a modified swallowing process adapted for rapid intake during flight or foraging, featuring a crop—a dilated esophageal pouch—for temporary food storage before propulsion into the proventriculus.^[74] Swallowing occurs via esophageal peristalsis, often aided by neck extension, which squeezes the bolus downward without the need for extensive oral manipulation.^[74] In the proventriculus, the glandular stomach secretes digestive enzymes to initiate breakdown, particularly effective for birds consuming whole prey like fish with bones.^[74] This storage and propulsion mechanism enhances feeding efficiency in aerial predators, such as raptors, by decoupling ingestion from immediate digestion.^[74] Amphibians and reptiles utilize gular pumping, involving expansion of the throat region primarily to assist in respiration, often combined with lingual retraction to draw food into the oral cavity.^[75] In monitor lizards, this positive-pressure gular pump ventilates lungs during locomotion. Frogs exemplify lingual retraction in feeding, where the tongue projects to capture prey and retracts to position it for swallowing, with subsequent pharyngeal compression propelling the bolus.^[76] Reptiles like lizards transport food intraorally using tongue cycles synchronized with hyoid and jaw movements, lacking the mammalian epiglottis and instead depending on glottal closure for airway protection. Invertebrate adaptations highlight further diversity, with insects relying on pharyngeal pumps for ingesting liquid diets through cibarial and pharyngeal musculature that creates suction.^[77] In blood-feeding bugs like Rhodnius prolixus, the pharyngeal pump adjusts stroke volume and frequency based on fluid viscosity, enabling efficient uptake of thin liquids without solid bolus formation.^[77] Cephalopods, such as octopuses, form boluses using beak-assisted biting and radula grinding, with arms manipulating prey pieces before esophageal swallowing.^[78] The chitinous beak shears food into manageable segments, which are then propelled by peristalsis, bypassing the need for extensive oral processing seen in vertebrates.^[78] Key differences from mammalian swallowing include the widespread absence of an epiglottis in non-mammals, reducing specialized laryngeal protection and emphasizing alternative safeguards like glottal adduction or positional anatomy.^[79] Some aquatic species, particularly certain fish and amphibians, incorporate ciliary action in the esophagus to aid bolus progression in low-viscosity environments, contrasting with the muscular peristalsis dominant in mammals.^[79] These adaptations reflect evolutionary trade-offs, such as integrating swallowing with gill ventilation in fish or aspiration breathing in reptiles.^[79] Swallowing efficiency in non-mammals often ties to ecological roles in predation, as seen in frogs where rapid tongue projection and retraction enable capture of evasive insects, enhancing survival in insect-rich habitats.^[76] This mechanism supports frogs as key predators in ecosystems, controlling pest populations while serving as prey for larger animals, thus maintaining trophic balance.^[76]

References

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Physiology, Swallowing - StatPearls - NCBI Bookshelf - NIH
Jul 24, 2023 · The process of swallowing, also known as deglutition, involves the movement of substances from the mouth (oral cavity) to the stomach via the pharynx and ...
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Anatomy and Physiology of Feeding and Swallowing – Normal ... - NIH
Eating and swallowing are complex behaviors involving volitional and reflexive activities of more than 30 nerves and muscles.
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Anatomy, Head and Neck, Swallowing - StatPearls - NCBI Bookshelf
Swallowing sounds like a simple physiological human function, but it is a complex, multifaceted process involving a variety of muscles and nerves.
[4]
[PDF] History of Oropharyngeal Organs and Swallowing
During swallowing, the airway must be closed to prevent swallowed material from entering the airway. Impairments to this coordination caused by dysphagia ...
[5]
Acorn worm genome reveals gill origins of human pharynx
Nov 19, 2015 · The newly sequenced genomes of two marine worms are shedding light on the 570-million-year evolution of gills into the pharynx that today gives humans the ...
[6]
How the tongue shaped life on Earth | Science | AAAS
May 25, 2023 · By making it possible to ingest food without suction, the evolution of the tongue some 350 million years ago was key to enabling vertebrates to ...
[7]
Evolution and Development of Dual Ingestion Systems in Mammals
Put simply, our thesis is that in every mammal there are at least two, separable ingestion systems, namely, the suckling system and the feeding system. Each ...
[8]
Mammals' weird way of swallowing is at least 165 million years old
Jul 18, 2019 · A new fossil find may help pinpoint the origins of mammals' uber-flexible hyoid bone, which anchors the tongue and gives us our signature swallowing style.
[9]
The Evolution and Development of Human Swallowing - Ento Key
Apr 1, 2017 · The permanent intersection of the respiratory and digestive pathways has created a de novo aerodigestive tract; a first of its kind in mammals.<|control11|><|separator|>
[10]
Vertebrate Evolution Conserves Hindbrain Circuits despite Diverse ...
Mar 11, 2021 · In this review, we summarize major modes of feeding and breathing and principles underlying their coordination in many vertebrate species.
[11]
[PDF] Comparative Chewing Efficiency in Mammalian Herbivores
Therefore, after a certain number of chews, swallowing the bolus will be more efficient than continuing to chew; this process intrinsically constrains the ...
[12]
Swallowing function and nutritional status in Japanese elderly ...
The findings indicate that maintaining swallowing function may contribute to the prevention of malnutrition in frail elderly people.
[13]
Swallow Function in Advanced Age
Age-related changes in swallow function are observed starting at approximately age 60; by age 80, most individuals will experience some degree of age-related ...<|control11|><|separator|>
[14]
Dysphagia in the elderly: management and nutritional considerations
Jul 30, 2012 · Of particular interest are recent studies that implicate benefit from intensive swallowing rehabilitation in preventing nutritional decline and ...
[15]
Anatomy, Head and Neck, Oral Cavity (Mouth) - StatPearls - NCBI
The teeth, which are the chief structures of the oral cavity, tear and grind ingested food into pieces small enough for digestion. The tongue enables the ...
[16]
Anatomy, Thorax, Esophagus - StatPearls - NCBI Bookshelf
The innervation of the esophagus involves the sympathetic and parasympathetic nervous systems, with primary innervation being sourced from the vagus nerve and ...
[17]
Chapter 53: THE PHARYNX AND LARYNX
Innervation and blood supply The motor and most of the sensory supply to the pharynx is by way of the pharyngeal plexus, which, situated chiefly on the middle ...
[18]
Brain stem control of swallowing: neuronal network and ... - PubMed
Swallowing movements are produced by a central pattern generator located in the medulla oblongata. It has been established on the basis of microelectrode ...
[19]
Brain stem control of the phases of swallowing - PubMed
Apr 28, 2009 · The nucleus tractus solitarius (NTS) probably contains the second-order sensory neurons as well as the pattern-generating circuitry of both the ...
[20]
Neurogenic Dysphagia: Peripheral and Central Neuromodulation
Sep 4, 2025 · Swallowing is regulated by a central pattern generator (CPG) located in the medulla oblongata. This center is bilaterally represented, and ...
[21]
Central Nervous System Control of Voice and Swallowing - PMC
Thus the same neurons are involved in central pattern generator for respiration and swallowing. ... Neuronal activity in the medulla oblongata during ...
[22]
Cortical and Subcortical Control of Swallowing—Can We Use ...
Jul 10, 2020 · The brainstem includes parts of the corticobulbar tracts with the cranial nerve nuclei that directly send and receive projections from muscles ...Missing: voluntary | Show results with:voluntary
[23]
Central Program Generator and Brain Stem - NCBI - NIH
Briefly, the critical elements of CPG are nucleus tractus solitarius (NTS) ... nucleus ambiguus and dorsomotor nucleus of vagus (Figure 1). All of these ...
[24]
Sensory Input Pathways and Mechanisms in Swallowing: A Review
Afferent input related to swallowing travels via sensory fibers in the trigeminal nerve (Vth), the glossopharyngeal nerve (IXth), the internal branch of the ...
[25]
The relationship between the oral and pharyngeal phases of ...
The oral phase of swallowing is a voluntary event, and the pharyngeal phase is an involuntary, independent event.1 However, swallowing occurs in a sequence ...
[26]
Coordination of Mastication, Swallowing and Breathing - PMC - NIH
... swallowing is essential to provide proper food nutrition and to prevent pulmonary aspiration. ... important for preventing aspiration of the liquid. When ...
[27]
Effects of aging on brain networks during swallowing - NIH
Jan 13, 2021 · These results suggest that aging has negative effects on the activation of swallowing-related regions and task-induced deactivation of the DMN.
[28]
Dysphagia in the Elderly - PMC - PubMed Central
We review age-related changes in peripheral and central nervous system control of head and neck structures for swallowing in this paper. In addition, we ...
[29]
https://pmc.ncbi.nlm.nih.gov/articles/PMC7806781/
[30]
https://pmc.ncbi.nlm.nih.gov/articles/PMC3182519/
[31]
Oral transit time: a critical review of the literature - PubMed Central
The times found varied from 0.35 s to 1.54 s for liquids, from 0.39 s to 1.05 s for pasty foods and from 1 s to 12.8 s for solid foods.
[32]
Anatomy and Physiology of Swallowing - Physiopedia
The oral propulsive phase immediately follows the oral preparatory phase: the tip of the tongue rises to touch the hard palate; as the tip of the tongue rises, ...Missing: variability | Show results with:variability
[33]
Adult Swallow Physiology – Swallowing and its Disorders Across the ...
The goal of the pharyngeal phase of the swallow is to transport the bolus through the pharyngeal conduit toward the esophagus without breaching the nasal cavity ...Oral Swallow Physiology · Pharyngeal Swallow... · Pharyngeal Pressure
[34]
Stages of swallowing: Deglutition - Kenhub
The food bolus is soft enough that it can easily be swallowed and propelled through the alimentary canal. Once food is swallowed, from the mouth it moves into ...Missing: variability texture
[35]
Evaluation and Treatment of Swallowing Impairments - AAFP
Apr 15, 2000 · Patients with poor pharyngeal contraction usually have more pharyngeal retention with thickened liquids and chewed solid foods than with thin ...
[36]
Physiology, Esophagus - StatPearls - NCBI Bookshelf
May 1, 2023 · The esophagus is a muscular channel that carries food from the pharynx to the stomach. It starts with the upper esophageal sphincter, formed in part by the ...
[37]
Physiology of oral, pharyngeal, and esophageal motility - Nature
May 16, 2006 · The neuromuscular structure and control of the oral, pharyngeal, and esophageal phases of swallowing are different.
[38]
Dysphagia: Practice Essentials, Background, Anatomy
May 7, 2024 · Additional iatrogenic causes of dysphagia. These include the following: Use of a cervical brace. Ventilator dependency. Psychogenic dysphagia.<|control11|><|separator|>
[39]
Dysphagia - StatPearls - NCBI Bookshelf - NIH
Nov 18, 2023 · Dysphagia, difficulty swallowing, can be acute or chronic and may occur in either the oropharyngeal or esophageal phases of swallowing.Missing: Hippocrates | Show results with:Hippocrates
[40]
Achalasia - Symptoms and causes - Mayo Clinic
Jul 19, 2024 · Symptoms · Difficulty swallowing, called dysphagia, which may feel like food or drink is stuck in the throat. · Swallowed food or saliva flowing ...
[41]
Globus Sensation (Lump in Throat) - Cleveland Clinic
Oct 15, 2024 · Globus sensation differs from dysphagia (difficulty swallowing) and odynophagia (painful swallowing). Globus sensation isn't painful. But ...
[42]
Aspiration from Dysphagia | Cedars-Sinai
About half of people with dysphagia have aspiration. About one-third of these people will need treatment for pneumonia at some point.
[43]
Preventing Aspiration in Older Adults with Dysphagia | HIGN
In fact, the risk of pneumonia is three times higher in patients with dysphagia (Hebert et al., 2016). Other harmful sequelae of dysphagia include malnutrition ...
[44]
Dysphagia Among Nursing Home Residents: An Assessment and ...
May 5, 2021 · The literature suggests that 40% to 60% of nursing home residents have some degree of dysphagia, i.e., difficulty in swallowing. Poorly managed, ...
[45]
The Prevalence of Dysphagia in Individuals Living in Residential ...
Mar 13, 2024 · Dysphagia prevalence ranged from 16 to 69.6%. The pooled prevalence of dysphagia was 56.11% (95% CI 39.363–72.172, p < 0.0001, I2 = 98.61%).
[46]
Oropharyngeal Dysphagia as the Main Expression of Amyotrophic ...
May 9, 2022 · ALS tends to manifest as limb weakness, but some patients present with bulbar symptoms, such as dysphagia and dysarthria.Missing: post- | Show results with:post-
[47]
Oropharyngeal dysphagia in older persons - PubMed Central - NIH
Older patients are particularly vulnerable to dysphagia because multiple age-related changes increase the risk of dysphagia. Physicians in charge of older ...
[48]
https://pmc.ncbi.nlm.nih.gov/articles/PMC9146888/
[49]
Videofluoroscopic Swallowing Study (VFSS) - Cleveland Clinic
Sep 2, 2025 · Videofluoroscopic swallow study (VFSS) is a medical term for the test that checks how you swallow specific food and liquids. It's also known as ...
[50]
Fiberoptic Evaluation of Swallowing | Johns Hopkins Medicine
A fiberoptic endoscopic evaluation of swallowing (FEES) test is a procedure used to assess how well you swallow. During the procedure, a speech-language ...
[51]
https://my.clevelandclinic.org/health/diagnostics/videofluoroscopic-swallowing-study-vfss
[52]
Esophageal manometry - Mayo Clinic
Jul 9, 2024 · This test can be helpful in diagnosing esophageal conditions, especially if you have trouble swallowing. The esophagus is a long, muscular ...
[53]
Esophageal Manometry | Johns Hopkins Medicine
Esophageal manometry helps determine what is causing these symptoms, which could be conditions such as: Achalasia (esophagus muscles don't help move food down ...
[54]
https://www.mayoclinic.org/tests-procedures/esophageal-manometry/about/pac-20394000
[55]
Effects of Mendelsohn Maneuver on Measures of Swallowing ...
A successful Mendelsohn swallow meant the participant was able to swallow and sustain laryngeal elevation for approximately 2 seconds or greater. Using SEMG for ...
[56]
Thickening agents used for dysphagia management: effect on ...
May 1, 2013 · Thickened liquids are often used in the management of dysphagia to improve bolus control and to help prevent aspiration.
[57]
Heller Myotomy: What It Treats, Surgery Steps & Recovery
Heller myotomy is a treatment for achalasia. It's a type of thoracic surgery that involves a surgeon making small cuts in your lower esophageal sphincter.
[58]
Peroral Endoscopic Myotomy (POEM) - Johns Hopkins Medicine
POEM can be a treatment option for people with muscle disorders in the esophagus, such as achalasia. POEM generally takes up to one hour and is performed ...
[59]
Swallowing therapy for dysphagia in acute and subacute stroke
Oct 30, 2018 · However, swallowing therapy may have reduced length of hospital stay, dysphagia, and chest infections, and may have improved swallowing ability.
[60]
Thick Liquids and Clinical Outcomes in Hospitalized Patients With ...
May 6, 2024 · A study of 92 patients with neurogenic dysphagia found lower rates of aspiration with mildly thick liquids compared with thin liquids (8.3% vs ...
[61]
Aspiration prevention surgeries: a review - Respiratory Research
Feb 6, 2023 · Furthermore, aspiration prevention surgeries improve the quality of life of patients and their caregivers by decreasing suctioning frequency.
[62]
Deep learning-based video analysis for automatically detecting ...
Jul 7, 2025 · Currently, the videofluoroscopic swallowing study (VFSS) is the gold standard for diagnosing dysphagia and assessing its severity. It ...
[63]
Device-based solutions supporting patients with swallowing problems
In summary, rTMS holds promise as a noninvasive neuromodulation technique for improving swallowing function in post-stroke dysphagia ( Table 1 ). Further ...
[64]
Artificial intelligence in dysphagia since the 21st century
Aug 18, 2025 · AI is driving significant shifts in both research and clinical practice in dysphagia; however, challenges such as interdisciplinary integration ...
[65]
Current perspectives on eating and rumination activity in dairy cows
Ruminants chew their feed initially during eating, and swallowed feed is later regurgitated and remasticated through the process of rumination.
[66]
Using rumination time to manage health and reproduction in dairy ...
Sep 26, 2021 · Regurgitation exposes animals to a reticular contraction, which along with the relaxation of the distal esophageal sphincter, allows a bolus of ...
[67]
Cutting food in terrestrial carnivores and herbivores - PMC - NIH
Insects and mammals cut their food up into small pieces to facilitate ingestion and chemical digestion. Teeth and jaws act as cutting tools.
[68]
Histological and histochemical characteristics of the esophagus in ...
Morphologically, the results revealed that the total esophageal length of the local breed donkey ranged from 89–110 cm, 200.32 ± 4.32 gm weight, and 1.54 ± 0.20 ...
[69]
XROMM and diceCT reveal a hydraulic mechanism of tongue base ...
May 19, 2020 · In primates, swallow jaw gape cycles consist of fast close, slow close, slow open, and fast open phases and are longer in duration than chewing ...Missing: mobility | Show results with:mobility
[70]
Videofluorographic assessment of deglutitive behaviors in a rat ...
Bolus Area, Size of the bolus (mm2) ; Swallow Rate, Following PB engagement, the number of swallows over 15 sec. This measure was not included among our final ...
[71]
Suckling, Feeding, and Swallowing: Behaviors, Circuits, and Targets ...
In infants, a suck-swallow-breathe cycle is in place at birth and then advances toward a chew-swallow-breathe cycle (Matsuo & Palmer 2015). To suck liquids, ...
[72]
Vibroacoustic Response of the Tympanic Membrane to Hyoid-Borne ...
The echolocation calls of bats are generated either via tongue clicking or by vibration of the vocal cords in the larynx. Laryngeal echolocation in bats ...
[73]
An XROMM Study of Food Transport and Swallowing in Channel ...
Prior work indicates that, in general, fish use the pharyngeal jaws to manipulate food into the esophagus, where peristalsis is thought to take over. We used X- ...
[74]
Digestive Anatomy and Physiology of Birds
Swallowing is accomplished by esophageal peristalsis, and in most birds appears to be aided by extension of the neck. Most but not all birds have a crop, which ...
[75]
Contribution of gular pumping to lung ventilation in monitor lizards
Evidence presented here shows that, during locomotion, varanids use a positive pressure gular pump to assist lung ventilation.
[76]
Prey Capture in Frogs: Alternative Strategies, Biomechanical Trade ...
Aug 10, 2025 · Frogs and toads capture insects using their sticky tongues (Fig. 1e). Then they take the insects into their mouths and swallow them (Monroy & ...
[77]
Effect of Diet Viscosity on the Operation of the Pharyngeal Pump in ...
Oct 1, 1979 · One of the striking anatomical features of the blood-feeding bug Rhodnius prolixus (Stahl) is the highly developed pharyngeal (or cibarial) pump ...
[78]
Feeding and Foraging (Chapter Four) - Cephalopod Behaviour
Mar 9, 2018 · This means that prey is not swallowed whole but is broken up by the combined actions of the beak ... Cephalopods feed mainly on molluscs, ...<|separator|>
[79]
Vertebrate Evolution Conserves Hindbrain Circuits despite Diverse ...
Reptiles and mammals have different swallowing procedures. In many reptile species like lizards and turtles, swallowing consists of two discrete stages.