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— CH. 1 · INTRODUCTION —

Cryptomonad

~4 min read · Ch. 1 of 5
5 sections
  • Cryptomonads are single-celled organisms so small that dozens could fit across the width of a human hair, yet they carry inside them one of the strangest evolutionary secrets in all of biology. Each cell runs between 10 and 50 micrometres long, flattened like a tiny leaf, with a groove or pocket near its front end and a pair of slightly unequal whip-like flagella beating at its rim. In freshwater ponds, in coastal seas, in brackish estuaries, these organisms are quietly everywhere. But what makes them remarkable is not their abundance. It is what lives inside them. Hidden between the inner membranes of their chloroplasts sits a shrunken cell nucleus called a nucleomorph, the remnant of an ancient red alga that was swallowed whole and never fully digested. How a eukaryote ends up inside another eukaryote, and what happens over deep time when it does, is the question that makes cryptomonads one of the most studied puzzles in the science of cell evolution.

  • Christian Gottfried Ehrenberg first noted cryptomonads in 1831 while studying Infusoria, though he would not have had the tools to see what was truly going on inside them. The cell body is wrapped not in an ordinary membrane but in a periplast, a cryptophyte-specific outer covering. Beneath it, and associated with the anterior pocket, sit structures called ejectisomes: two connected spiral ribbons held under tension like a coiled spring. When the cell is disturbed by mechanical force, chemical stress, or a sudden change in light, those ribbons fire. The discharge propels the cell away in a zig-zag course, a microscopic escape manoeuvre. Larger ejectisomes sit near the pocket and are visible under a standard light microscope; smaller ones are distributed beneath the periplast across the cell body. The flagella themselves are not bare whips. They are covered in bipartite hairs called mastigonemes, assembled inside the endoplasmic reticulum and carried to the surface, with small scales sometimes present on both the flagella and the cell body.

  • Most cryptomonads carry one or two chloroplasts that can appear brown, red, or bluish-green depending on their pigment mix. Each chloroplast is wrapped in four membranes rather than the two found in most algae, and between the middle two membranes sits the nucleomorph. Genetic studies confirmed what the structure implied: the plastid originally belonged to a red alga that was engulfed by an ancestral cryptomonad host in a process called secondary endosymbiosis. The plastids, however, are strikingly different from those of living red algae. Phycobiliproteins are present, but only inside the thylakoid lumen, and only in the forms phycoerythrin or phycocyanin. In the genus Rhodomonas, researchers have resolved the crystal structure of a phycobiliprotein to 1.63 angstroms, and found that the alpha subunit bears no relation to any other known phycobiliprotein. Meanwhile, Cryptomonas paramecium, once called Chilomonas paramecium, has leucoplasts rather than photosynthetic chloroplasts, and the class Goniomonadea lacks plastids entirely, an arrangement that supports the view that the cryptomonad lineage was originally heterotrophic and only later acquired the ability to photosynthesize.

  • Some Cryptomonas species can shift into a palmelloid stage, clustering together inside a shared mucus, only to dissolve back into free-living flagellates when conditions improve. Certain species go further still, forming immotile cysts with rigid cell walls to ride out periods of stress. Mitosis in cryptomonads is open, meaning the nuclear envelope breaks down during cell division. Sexual reproduction has also been documented, though it is not universal across the group. A particularly striking strategy is mixotrophy: some cryptomonads can both photosynthesize and consume organic matter, giving them flexibility when light or nutrients are scarce. The flat cristae of their mitochondria link them structurally to other groups in the tree of life, a detail that has been central to debates about how the major algal lineages relate to one another.

  • Botanists historically placed cryptomonads in the class Cryptophyceae or the division Cryptophyta; zoologists treated the same organisms as the flagellate protozoan order Cryptomonadina. For a time, their pigmentation looked similar enough to that of dinoflagellates that some classifications grouped both lineages together as the Pyrrhophyta. Cavalier-Smith later united cryptomonads with the heterokonts and haptophytes as the Chromista, based on the similarity of their chloroplasts. That proposal ran into trouble when ultrastructural analysis revealed deep differences in overall cell organisation among the three groups, suggesting the three major chromist lineages had each acquired plastids independently and that the grouping was polyphyletic. Molecular work by Parfrey and colleagues, and separately by Burki and colleagues, placed Cryptophyceae instead as a sister clade to the green algae, or to green algae combined with glaucophytes. The sister group to the entire cryptomonad superclass is likely the kathablepharids, flagellates that share the ejecting ejectisomes but do not have plastids. Genetic studies dating back to 1994 supported the placement of Goniomonas as sister to Cryptophyceae within the superclass, helping anchor what had long been a structurally defined grouping onto a molecular foundation.

Common questions

What are cryptomonads and where are they found?

Cryptomonads, also called cryptophytes, are a superclass of single-celled algae and colorless flagellates. Each cell is around 10-50 micrometres in size. They are common in freshwater habitats and also occur in marine and brackish environments.

Why do cryptomonads have four membranes around their chloroplasts?

Cryptomonad chloroplasts are surrounded by four membranes because they were acquired through secondary endosymbiosis: an ancestral host engulfed a red alga, whose own double membrane was retained alongside the host's membrane layers. A remnant nucleus called the nucleomorph sits between the inner two membranes.

What is a cryptomonad nucleomorph?

The nucleomorph is a reduced cell nucleus found between the middle two membranes of the cryptomonad chloroplast. It is the vestigial nucleus of the red alga that was engulfed during secondary endosymbiosis, confirmed by genetic studies.

What are ejectisomes in cryptomonads?

Ejectisomes are coiled extrusomes made of two connected spiral ribbons held under tension. When a cryptomonad is disturbed by mechanical, chemical, or light stress, the ejectisomes discharge and propel the cell away in a zig-zag course.

Who first described cryptomonads and when?

Christian Gottfried Ehrenberg made the first recorded mention of cryptomonads in 1831 while studying Infusoria. Later, botanists classified them as the algae division Cryptophyta while zoologists placed them in the protozoan order Cryptomonadina.

What is the sister group to cryptomonads?

The sister group to the cryptomonads is most likely the kathablepharids, a group of flagellates that also possess ejectisomes. Molecular studies by Parfrey and Burki placed Cryptophyceae as a sister clade to the green algae, or to green algae plus glaucophytes.

All sources

17 references cited across the entry

  1. 1journalKingdom Chromista and its eight phyla: A new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergencesThomas Cavalier-Smith — 2018
  2. 2journalThe Katablepharids are a Distant Sister Group of the Cryptophyta: A Proposal for Katablepharidophyta Divisio Nova/Kathablepharida Phylum Novum Based on SSU rDNA and Beta-Tubulin PhylogenyN. Okamoto et al. — 2005
  3. 4bookOrganelles, Genomes, and Eukaryote PhylogenyThomas Cavalier-Smith — 2004
  4. 5citationGoniomonas: rRNA sequences indicate that this phagotrophic flagellate is a close relative of the host component of cryptomonadsMcFadden, Gilson, & Hill — 1994
  5. 6bookAlgaeL. E. Graham et al. — Benjamin Cummings (Pearson) — 2009
  6. 7journalA comparison of the periodic sub-structures of the trichocysts of the Cryptophyceae and PrasinophyceaeS. Morrall et al. — 1980
  7. 8journalThe ejectisomes of the flagellate Chilomonas paramecium - Visualization by freeze-fracture and isolation techniquesJ. N. Grim et al. — 1984
  8. 10journalThe highly reduced genome of an enslaved algal nucleusS. Douglas — 2002
  9. 11journalEvolution of a light-harvesting protein by addition of new subunits and rearrangement of conserved elements: Crystal structure of a cryptophyte phycoerythrin at 1.63Å resolution.K. Wilk — 1999
  10. 13journalNuclear genome sequence of the plastid-lacking cryptomonad Goniomonas avonlea provides insights into the evolution of secondary plastidsU. Cenci et al. — 2018
  11. 14journalEstimating the timing of early eukaryotic diversification with multigene molecular clocksLaura Wegener Parfrey et al. — August 16, 2011
  12. 15journalUntangling the early diversification of eukaryotes: a phylogenomic study of the evolutionary origins of Centrohelida, Haptophyta and CryptistaFabien Burki et al. — 2016-01-27
  13. 16journalMitochondrial Genomes of Hemiarma marina and Leucocryptos marina Revised the Evolution of Cytochrome c Maturation in CryptistaYuki Nishimura et al. — 2020