Morphallaxis

Morphallaxis is the regeneration of specific tissue in a variety of organisms due to loss or death of the existing tissue. The word comes from the Greek allazein, (αλλάζειν) which means to change.

The classical example of morphallaxis is that of the Cnidarian hydra, where when the animal is severed in two (by actively cutting it with, for example, a surgical knife) the remaining severed sections form two fully functional and independent hydra. The notable feature of morphallaxis is that a large majority of regenerated tissue comes from already-present tissue in the organism. That is, the one severed section of the hydra forms into a smaller version of the original hydra, approximately the same size as the severed section. Hence, there is an "exchange" of tissue.

Researchers Wilson and Child showed circa 1930 that if the hydra was pulped and the disassociated food passed through a sieve, those cells then put into an aqueous solution would shortly reform into the original organism with all differentiated tissue correctly arranged.[1]

Morphallaxis is often contrasted with epimorphosis, which is characterized by a much greater relative degree of cellular proliferation. Although cellular differentiation is active in both processes, in morphallaxis the majority of the regeneration comes from reorganization or exchange, while in epimorphosis the majority of the regeneration comes from cellular differentiation. Thus, the two may be distinguished as a measure of degree. Epimorphosis is the regeneration of a part of an organism by proliferation at the cut surface. For example, in Planaria neoblasts help in regeneration.

History

The word comes from the Greek allazein, which means to exchange.[1] The biological process was first discovered in hydra by Abraham Trembley, who was considered the father of environmental zoology. Abraham Trembley was doing a research on a sample pond water and examined the lifestyle of hydra. He couldn’t decide if they belong to the animal or plant kingdom, so he cut them in half and planned to see whether they die, like animals would, or re-pattern, as plants. Even though the halves and smaller pieces gave rise to new individuals, he still believed that hydra is an animal, since all their features, like movements or feeding behavior matched with animals’. Trembley came to the conclusion, that some animals have the ability to regenerate.[2]

The process and mechanism of planarian regeneration was eventually renamed to 'Morphallaxis' by Thomas Hunt Morgan, the father of experimental genetics.[2]

Regeneration in Cnidarian hydras

Hydras are a group of freshwater Cnidarians that are about 0.5 cm long. A hydra has a short, tubular shaped body. Hydras have a head that consists of a hypostome region and a foot that consists of a basal disc. The head portion of the hydra contains the mouth and tentacles, which allows for the catching and eating of food. The foot portion of the hydra contains the basal disc which allows for the hydra to stick to rocks and other elements.[3]

When a hydra gets cut in half, the head portion can regenerate and form a new foot with the basal disc, as well as, the foot portion can regenerate and form a new head with the hypostome region. If a hydra was severed into smaller pieces, the middle pieces would still form a head and foot at the appropriate regions of the hydra. This results in a smaller hydra that was regenerated by morphallaxis and occurs without cellular division.[3]

Mechanism

The mechanism involved uses regenerative tissue remodeling. This allows the body's axes to repattern and form new tissue, as well as causes organs in the body to redevelop into different proportions.

Hydras contain a series of gradients that controls the formation of the correct head and foot regeneration. The head gradient permits the head to only form in one place and the foot gradient permits the basal disc to only form in another place. These gradients are driven by the polarity in the hydra. The hypostome in the head region inhibits the formation of another hypostome. This explains why two heads will not form on one hydra.

Types of Regeneration

There are three types of regenerations. One of them is Epimorphosis, which appears in Salamander limbs. This type of regeneration includes the formation of a new part, which is called the blastema. The amputation is sensed by a large number of somatic stem cells, that migrate to the wound they increase their division rate. At the wound blastema forms and the blastema cells proliferate to re-generate the lost tissues. There is no significant re-patterning of the remaining tissue.[4][5]

Another type of regeneration is Morphallaxis, which is usually observed in Hydras. The main difference between the two types is that morphallatic regeneration does not include the formation of blastemal and there is no proliferation. Instead the existing tissue undergoes re-arrangement and it is transformed into the new organ.[4][5]

The third type occurs for example in planarians. It was discovered once regeneration was observed in a cellular level. For very long biologists believed that planarians undergo epimorphosis, because regeneration on a trunk piece shows new tissue formation from a blastemal. However, on a tail piece next to the formation of a blastemal (which serves as a signaling center in this case), pharynx was re-arranged from a pre-existing tissue. The conclusion is that planarian regeneration cannot be listed neither to Epimorphosis or Morphallaxis.[5]

References

1. Dunbar HF (1946). Emotions and Bodily Changes. New York.
2. Pellettieri J (March 2019). "Regenerative tissue remodeling in planarians - The mysteries of morphallaxis". Seminars in Cell & Developmental Biology. 87: 13–21. doi:10.1016/j.semcdb.2018.04.004. PMC 6195476. PMID 29631028.
3. Gilbert SF (2000). "Regeneration". Developmental Biology (6th ed.). ISBN 978-0878932436.
4. Pellettieri, Jason (March 2019). "Regenerative tissue remodeling in planarians – The mysteries of morphallaxis". Seminars in Cell & Developmental Biology. 87: 13–21. doi:10.1016/j.semcdb.2018.04.004. ISSN 1084-9521. PMC 6195476. PMID 29631028.
5. Agata, Kiyokazu; Saito, Yumi; Nakajima, Elizabeth (2007-02-28). "Unifying principles of regeneration I: Epimorphosis versus morphallaxis". Development, Growth & Differentiation. 49 (2): 73–78. doi:10.1111/j.1440-169x.2007.00919.x. ISSN 0012-1592. PMID 17335428.