Protist locomotion

Motion system of a type of eukaryotic organism

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amoeboids. Other protists are not motile

, and consequently have no built-in movement mechanism.

Overview

Unicellular protists comprise a vast, diverse group of organisms that covers virtually all environments and habitats, displaying a menagerie of shapes and forms. Hundreds of species of the ciliate genus Paramecium ^[3] or flagellated Euglena ^[4] are found in marine, brackish, and freshwater reservoirs; the green algae Chlamydomonas is distributed in soil and fresh water world-wide;^[5] parasites from the genus Giardia colonize intestines of several vertebrate species.^[6] One of the shared features of these organisms is their motility, crucial for nutrient acquisition and avoidance of danger.^[7] In the process of evolution, single-celled organisms have developed in a variety of directions, and thus their rich morphology results in a large spectrum of swimming modes.^[8][2]

Many swimming protists actuate tail-like appendages called

dynein motors, powered by ATP hydrolysis, perform mechanical work by promoting the relative sliding of filaments, resulting in bending deformations.^[2]

Although protist flagella have a diversity of forms and functions,[11] two large families, flagellates and ciliates, can be distinguished by the shape and beating pattern of their flagella.^[2]

In the phylogenetic tree on the right, aquatic organisms (living in marine, brackish, or freshwater environments) have their branches drawn in blue while parasitic organisms have their branches drawn in red. Ciliates are indicated by an asterisk after their names. For each phylum marked in bold font, a representative organism has been sketched next to its name.^[2]

Modes of locomotion

Protists according to how they move
Type of protist		Movement mechanism	Description	Example		Other examples
Motile	Flagellates		A flagellum (Latin for whip) is a lash-like appendage that protrudes from the cell body of some protists (as well as some bacteria). Flagellates use from one to several flagella for locomotion and sometimes as feeding and sensory organelle.		Cryptophytes	All foraminiferans and Apicomplexa )
	Ciliates		A cilium (Latin for eyelash) is a tiny flagellum. Ciliates use multiple cilia, which can number in many hundreds, to power themselves through the water.		Paramecium bursaria click to see cilia	amoebae, ciliates and flagellates .
	Amoebas (amoeboids)		Pseudopods (Greek for false feet) are lobe-like appendages which amoebas use to anchor to a solid surface and pull themselves forward. They can change their shape by extending and retracting these pseudopods.[14]		Amoeba	Found in every major protist fungi.[15][16]
Not motile		none			Diatom	Coccolithophores, most diatoms, and non‐motile species of Phaeocystis[13] Among protozoans the parasitic Apicomplexa are non‐motile.

Flagellates

Flagella

cilia

are widely used in eukaryotic cells (plant and animal) apart from protists.

The regular beat patterns of eukaryotic cilia and flagella generates motion on a cellular level. Examples range from the propulsion of single cells such as the swimming of

spermatozoa to the transport of fluid along a stationary layer of cells such as in a respiratory tract

. Though eukaryotic flagella and motile cilia are ultrastructurally identical, the beating pattern of the two organelles can be different. In the case of flagella, the motion is often planar and wave-like, whereas the motile cilia often perform a more complicated three-dimensional motion with a power and recovery stroke.

Eukaryotic flagella—those of animal, plant, and protist cells—are complex cellular projections that lash back and forth. Eukaryotic flagella are classed along with eukaryotic motile cilia as undulipodia^[17] to emphasize their distinctive wavy appendage role in cellular function or motility. Primary cilia are immotile, and are not undulipodia.

Cryptaulax, Abollifer, Bodo, Rhynchomonas, Kittoksia, Allas, and Metromonas ^[18]

• 
  Freshwater green algal flagellate (Chlamydomonas)

Flagellates typically have a small number of long flagella distributed along the bodies, and they actuate them to generate thrust. The set of observed movement sequences includes planar undulatory waves and traveling helical waves, either from the base to the tip, or in the opposite direction.^[19][20] Flagella attached to the same body might follow different beating patterns, leading to a complex locomotion strategy that often relies also on the resistance the cell body poses to the fluid.^[2]

Ciliates

In contrast to flagellates, propulsion of ciliates derives from the motion of a layer of densely-packed and collectively-moving cilia, which are short hair-like flagella covering their bodies. The seminal review paper of Brennen and Winet (1977) lists a few examples from both groups, highlighting their shape, beat form, geometric characteristics and swimming properties.^[20] Cilia may also be used for transport of the surrounding fluid, and their cooperativity can lead to directed flow generation. In higher organisms this can be crucial for internal transport processes, as in cytoplasmic streaming within plant cells,^[21] or the transport of ova from the ovary to the uterus in female mammals.^[22][2]

Ciliates generally have hundreds to thousands of cilia that are densely packed together in arrays. Like the flagella, the cilia are powered by specialised molecular motors. An efficient forward stroke is made with a stiffened flagellum, followed by an inefficient backward stroke made with a relaxed flagellum. During movement, an individual cilium deforms as it uses the high-friction power strokes and the low-friction recovery strokes. Since there are multiple cilia packed together on an individual organism, they display collective behaviour in a metachronal rhythm. This means the deformation of one cilium is in phase with the deformation of its neighbor, causing deformation waves that propagate along the surface of the organism. These propagating waves of cilia are what allow the organism to use the cilia in a coordinated manner to move. A typical example of a ciliated microorganism is the Paramecium, a one-celled, ciliated protozoan covered by thousands of cilia. The cilia beating together allow the Paramecium to propel through the water at speeds of 500 micrometers per second.^[23]

• 
  Paramecium feeding on bacteria
• 
  The ciliate

  Oxytricha trifallax

  with cilia clearly visible
• Animation of swimming ciliate ^[24]

External videos
Paramecium: The white rat of ciliates
Flinching saves lives in the microcosmos – Journey to the Microcosmos
Microbes in slow motion – Journey to the Microcosmos

Amoebas

Locomotion by pseudopod

Amoeba proteus with many pseudopodia

See also: Amoeboid movement and Pseudopodia

The third prevalent forms of protist cell motility is actin-dependent cell migration. The evolution of flagellar-based swimming has been well studied, and strong evidence suggests a single evolutionary origin for the eukaryotic flagellum occurred before the diversification of modern eukaryotes. On the other hand, actin-dependent crawling uses many different molecular mechanisms, and the study of how these evolved is only just beginning.^[25]

Colonial protists

Gonium pectorale ^[26]

(a) Sixteen-cell colony. Each cell has two flagella, 30–40 μm long. Scale bar is 10 μm.
(b) Schematic of a colony of radius a: sixteen cells (green) each with one eye spot (orange dot). The cis flagellum is closest to the eye spot, the trans flagellum is furthest.^[27]

Gonium is a genus of colonial algae belonging to the family Volvocaceae. Typical colonies have 4 to 16 cells, all the same size, arranged in a flat plate, with no anterior-posterior differentiation. In a colony of 16 cells, four are in the center, and the other 12 are on the four sides, three each.^[28]

Since the work of August Weismann on germ-plasm theory in biology ^[29] and of Julian Huxley on the nature of the individual in evolutionary theory,^[30] the various species of green algae belonging to the family Volvocaceae have been recognized as important ones in the study of evolutionary transitions from uni- to multicellular life. In a modern biological view,^[31] this significance arises from a number of specific features of these algae, including the fact that they are an extant family (obviating the need to study microfossils), are readily obtainable in nature, have been studied from a variety of perspectives (biochemical, developmental, genetic), and have had significant ecological studies. From a fluid dynamical perspective,^[32] their relatively large size and easy culturing conditions allow for precise studies of their motility, the flows they create with their flagella, and interactions between organisms, while their high degree of symmetry simplifies theoretical descriptions of those same phenomena.^[33][26]

As they are photosynthetic, the ability of these algae to execute phototaxis is central to their life. Because the lineage spans from unicellular to large colonial forms, it can be used to study the evolution of multicellular coordination of motility. Motility and phototaxis of motile green algae have been the subjects of an extensive literature in recent years,^{[34][35][36][37][38][39][40]} focusing primarily on the two extreme cases: unicellular Chlamydomonas and much larger Volvox, with species composed of 1000–50,000 cells. Chlamydomonas swims typically by actuation of its two flagella in a breast stroke, combining propulsion and slow body rotation. It possesses an eyespot, a small area highly sensitive to light,^[41][42] which triggers the two flagella differently.^[43] Those responses are adaptive, on a timescale matched to the rotational period of the cell body,^[44][45][46] and allow cells to scan the environment and swim toward light.^[47] Multicellular Volvox shows a higher level of complexity, with differentiation between interior germ cells and somatic cells dedicated to propulsion. Despite lacking a central nervous system to coordinate its cells, Volvox exhibits accurate phototaxis. This is also achieved by an adaptive response to changing light levels, with a response time tuned to the colony rotation period which creates a differential response between the light and dark sides of the spheroid.^[35][48][26]

In light of the above, a natural questions is as follows: How does the simplest differentiated organism achieve phototaxis? In the Volvocine lineage the species of interest is Gonium. This 8- or 16-cell colony represents one of the first steps to true multicellularity,^[49] presumed to have evolved from the unicellular common ancestor earlier than other Volvocine algae.^[50] It is also the first to show cell differentiation.^[26]

The 16-cell Gonium colony shown in the diagram on the right is organized into two concentric squares of respectively 4 and 12 cells, each biflagellated, held together by an extracellular matrix.^[51] All flagella point out on the same side: It exhibits a much lower symmetry than Volvox, lacking anterior-posterior symmetry. Yet it performs similar functions to its unicellular and large colonies counterparts as it mixes propulsion and body rotation and swims efficiently toward light.^[34][52][53] The flagellar organization of inner and peripheral cells deeply differs:^[54][55] Central cells are similar to Chlamydomonas, with the two flagella beating in an opposing breast stroke, and contribute mostly to the forward propulsion of the colony. Cells at the periphery, however, have flagella beating in parallel, in a fashion close to Volvox cells.^[27] This minimizes steric interactions and avoids flagella crossing each other.^[34] Moreover, these flagella are implanted with a slight angle and organized in a pinwheel fashion [see Fig. 1(b)]:^[54] Their beating induces a left-handed rotation of the colony, highlighted in Figs. 1(c) and 1(d) and in Supplemental Movie 1 [29]. Therefore, the flagella structure of Gonium reinforces its key position as intermediate in the evolution toward multicellularity and cell differentiation.^[26]

These small flat assemblies show intriguing swimming along helical trajectories—with their body plane almost normal to the swimming direction—that have attracted the attention of naturalists since the 18th century.^[52][53][56] Yet the way in which Gonium colonies bias their swimming toward the light remains unclear. Early microscopic observations have identified differential flagellar activity between the illuminated and the shaded sides of the colony as the source of phototactic reorientation.^[52][53] Yet a full fluid-dynamics description, quantitatively linking the flagellar response to light variations and the hydrodynamic forces and torques acting on the colony, is still lacking. From an evolutionary perspective, phototaxis in Gonium raises fundamental issues such as the extent to which the phototactic strategy of the unicellular ancestor is retained in the colonial form, how the phototactic flagella reaction adapted to the geometry and symmetry of the colony, and how it leads to effective reorientation.^[26]

Protist taxis: Directed motion

Phototaxis