Spiracle (arthropods)

A spiracle or stigma is the opening in the exoskeletons of insects, myriapods, velvet worms and many arachnids to allow air to enter the trachea.[1][2][3] In insects, gas exchange occurs largely through a network of tracheae and air sacs that deliver oxygen directly to tissues and remove carbon dioxide; the circulatory system plays a comparatively minor role in oxygen transport and carbon dioxide removal. Tracheal tubes connect to the outside through spiracles, which can be specialized to regulate airflow and limit water loss.

Morphology and valve architecture

In insects, the external spiracular opening commonly leads into an atrium and a valve apparatus that regulates airflow into the tracheal trunks. The rim of the opening (often termed the peritreme) may bear cuticular hairs or filtering structures that reduce entry of dust, pathogens, and liquid water while also limiting convective water loss.[4] Comparative studies highlight substantial diversity in spiracle form across insect orders, including differences in atrial depth, cuticular filter design, and the mechanical arrangement of valve elements.[5] These anatomical features can be interpreted as tradeoffs among protection, resistance to airflow, and the speed and precision of opening and closing.

Control of spiracles

In most species, the spiracles are controlled by motor neurons in the central nervous system. It can be opened and closed in an efficient manner to admit air while minimizing associated physiological costs, such as water loss during respiration. Many sensory stimuli can affect the control of spiracles in insects, e.g. chemosensory (carbon dioxide, oxygen, etc.) or mechanosensory (sound, touch, etc.). It has been shown that during metabolically intensive behaviors, such as flight, insects can dynamically modulate the spiracle opening size just so to meet the metabolic demand of flight, while not losing too much water.[6]

Control of spiracle opening is done by a wide range of mechanisms, such as elastic closure, and closer muscles surrounding the spiracle or kinking the tube. In some the muscle relaxes to open the spiracle, in others to close it. [7] Several aquatic insects have similar or alternative closing methods to prevent water from entering the trachea. The timing and duration of spiracle closures can affect the respiratory rates of the organism.[8] Spiracles may also be surrounded by hairs to minimize bulk air movement around the opening, and thus minimize water loss.

In larger insects, spiracle control is more complex and critical for managing gas exchange due to their higher metabolic demands. Larger insects, such as locusts and some beetles, exhibit active ventilation, where spiracle control works in concert with abdominal movements. These abdominal contractions force air in and out of the tracheal system, and the spiracles open and close in a synchronized manner to maximize oxygen intake and carbon dioxide expulsion. This active process allows these insects to regulate their internal environment more precisely, especially during periods of high activity, such as flight. Research has shown that neural circuits in the insect's central nervous system adjust the spiracle opening in response to carbon dioxide concentration, ensuring efficient gas exchange and preventing hypoxia or hypercapnia. Other body parts, such as the proboscis,[9] might also extend or contract so as to ventilate the insect during various behaviors.

Spiracles in other animals besides insects

Most myriapods have paired lateral spiracles similar to those of insects. Scutigeromorph centipedes are an exception, having unpaired, non-closable spiracles at the posterior edges of tergites.[2]

Velvet worms have tiny spiracles scattered over the surface of the body and linked to unbranched tracheae. There can be as many as 75 spiracles on a body segment. They are most abundant on the dorsal surface. They cannot be closed, which means velvet worms easily lose water and thus are restricted to living in humid habitats.[3]

Although all insects have spiracles, only some arachnids have them. Some spiders such as orb weavers and wolf spiders have spiracles. Ancestrally, spiders have book lungs, not trachea. However, some spiders evolved a tracheal system independently of the tracheal system in insects, which includes independent evolution of the spiracles as well. These spiders retained their book lungs, however, so they have both.[10][11] Harvestmen, camel spiders, ricinuleids, mites, and pseudoscorpions all breathe through a tracheal system and lack book lungs.

Literature

  • Chapman, R.F. (1998): The Insects, Cambridge University Press

References

  1. ^ Solomon, Eldra, Linda Berg, Diana Martin (2002): Biology. Brooks/Cole
  2. ^ a b Hilken, Gero; Rosenberg, Jörg; Edgecombe, Gregory D.; Blüml, Valentin; Hammel, Jörg U.; Hasenberg, Anja; Sombke, Andy (2021). "The tracheal system of scutigeromorph centipedes and the evolution of respiratory systems of myriapods". Arthropod Structure & Development. 60 101006. Bibcode:2021ArtSD..6001006H. doi:10.1016/j.asd.2020.101006. PMID 33246291. S2CID 227191511.
  3. ^ a b "Untitled 1". lanwebs.lander.edu. Retrieved 6 February 2023.
  4. ^ Schmitz, A; Wasserthal, L. T (1 April 1999). "Comparative morphology of the spiracles of the Papilionidae, Sphingidae, and Saturniidae (Insecta: Lepidoptera)". International Journal of Insect Morphology and Embryology. 28 (1): 13–26. doi:10.1016/S0020-7322(98)00033-6. ISSN 0020-7322.
  5. ^ Hassan, A. A. G. (1944-07). "THE STRUCTURE AND MECHANISM OF THE SPIRACULAR REGULATORY APPARATUS I N ADULT DIPTERA AND CERTAIN OTHER GROUPS OF INSECTS". Transactions of the Royal Entomological Society of London. 94 (1): 103–153. doi:10.1111/j.1365-2311.1944.tb01214.x. ISSN 0035-8894. {{cite journal}}: Check date values in: |date= (help)
  6. ^ Lehmann, Fritz-Olaf (2001). "Matching Spiracle Opening to Metabolic Need During Flight in Drosophila". Science. 294 (5548): 1926–1929. Bibcode:2001Sci...294.1926L. doi:10.1126/science.1064821. PMID 11729318.
  7. ^ Imms' General Textbook of Entomology: Volume 1: Structure, Physiology and Development Volume 2: Classification and Biology. Berlin: Springer. 1977. ISBN 0-412-61390-5.
  8. ^ Wilmer, Pat, Graham Stone, and Ian Johnston (2005). Environmental Physiology of Animals. United Kingdom: Blackwell Publishing. pp. 171–172. ISBN 978-1-4051-0724-2.{{cite book}}: CS1 maint: multiple names: authors list (link)
  9. ^ Lehmann, Fritz-Olaf; Heymann, Nicole (2005). "Unconventional mechanisms control cyclic respiratory gas release in flying Drosophila". Journal of Experimental Biology. 208 (19): 3645–3654. Bibcode:2005JExpB.208.3645L. doi:10.1242/jeb.01788. PMID 16169942.
  10. ^ "How Do Spiders Breathe?". Sciencing. Retrieved 6 June 2021.
  11. ^ Schmitz, Anke (May 2016). "Respiration in spiders (Araneae)". Journal of Comparative Physiology B: Biochemical, Systemic, and Environmental Physiology. 186 (4): 403–415. doi:10.1007/s00360-016-0962-8. ISSN 1432-136X. PMID 26820263. S2CID 16863495.
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