I thought there were only two centrioles per cell, that convert to the basal body at some point during the cell cycle. I also thought there's one basal body per cilium, so I'm not clear on where the other basal bodies are coming from. I'd like to know the distribution of basal bodies and centrioles throughout the cell cycle in multi-ciliated organisms/cells.
There is a single basal body per cilium. During cell division the centrosome has two centrioles, however, during the differentiation of citiated cells, there is an amplification of basal bodies that nucleate from the centrioles.
In multiciliated cells the basal bodies arise from two pathways- 1. de-novo / deuterosomal / acentriolar pathway and 2. centriolar pathway.
In a work carried out by Al Jord et al (2014), the interplay of these two pathways had been studied in the brain ependymal cells. The study shows that post division, when the cell begins to differentiate, the daughter centriole serves as a site for creation of deuterosomes. During this stage deuterosome independent procentrioles also nucleate (the centriolar pathway) on both mother and daughter centrioles. However, the contribution to basal bodies is predominantly from the deuterosomes.
more than 90% of the centrioles were generated via deuterosomes and less than 10% directly from centrosomal centriole
Following figure explains the process nicely.
I am not really sure about multiciliated unicellular organisms. My guess is that they retain old basal bodies and new ones are created by a similar mechanism. Most ciliates can also undergo sexual reproduction.
It seems that there is one basal body per cilium in multiciliated cells. Centriole duplication (as far as I understand, a basal body is just a different name for a centriole that is attached to a cilium) is tightly coupled to the cell-cycle. There is always one pair of centrioles that are at the base of the primary cilium (of which there is exactly one in most cells).
But not in multiciliated cells. These cells are differentiated and can no longer divide, and their centriole duplication process is decoupled from the cell cycle. In fact, they aren't, strictly speaking, "duplicating", since many procentriolar bodies may form around one cylinder. Indeed, each cilium has its own basal body; thus a cell with hundreds of cilia has hundreds of basal bodies.
- Ishikawa 2011 is a review that examines the structure of primary and motile cilia, and ciliogenesis. It does not spell out how the basal body gets there.
- Anderson 1971 describes the formation of new cilia in monkey oviduct. They first remove the ovaries of these monkeys to prevent cells from differentiating (this apparently requires estrogen), then administer estrogen and look at EM images of the cells over the following 6 days. They describe at length the processes of basal body formation.
Dirksen 1991 is another review focusing specifically on basal body formation during ciliogenesis. From their introduction:
In order for the 200-300 cilia to form that the fully differentiated cell will need, production of that many centrioles is required first. The cell, after the last division, has usually retained a pair of centrioles. To what extent these mature centrioles are involved in the earlier stages of centriole formation, in which centriole precursor structures of varying sizes and shapes are generated and proceed to undergo transformational changes of great complexity, is largely unknown..
Sorokin 1968 is apparently a classic text which goes over EM images of ciliogenesis at various stages, and proposes a series of events describing what is happening. This is probably what you want, re: where are the other basal bodies coming from.
- Bettencourt-Dias 2007 reviews the research on the formation of new basal bodies. This is another publication that answers your question very directly:
Many ciliated cells, such as those in vertebrate respiratory and reproductive tracts, can have 200–300 cilia per cell. This requires the generation of multiple centrioles during ciliogenesis. Here, centrioles are generated by two pathways, a centriolar and an acentriolar mechanism. In the first one, a parent centriole usually produces one daughter at a time; however, in certain cases several centrioles have been observed to develop simultaneously around one parent centriole, with the daughter centrioles being released into the cytoplasm to mature. The acentriolar pathway is the major pathway for basal body production. In this pathway, fibrous granules 124 that are 70–100 nm in diameter first appear in the cytoplasm and subse quently move to the apical cytoplasm. Deuterosomes appear within the area. (...) Multiple procentrioles can grow out from deuterosomes, and mature daughter centrioles travel towards the apical region where they form ciliary basal bodies.