Diatom

Diatom
A diatom (Neo-Latin diatoma)[a] is any member of a large group

comprising several genera of algae, specifically microalgae, found in
the oceans, waterways and soils of the world. Living diatoms make
up a significant portion of the Earth's biomass: they generate about
20 to 50 percent of the oxygen produced on the planet each
year,[10][11] take in over 6.7 billion tonnes of silicon each year from
the waters in which they live,[12] and constitute nearly half of the
organic material found in the oceans. The shells of dead diatoms
can reach as much as a half-mile (800 m) deep on the ocean floor,
and the entire Amazon basin is fertilized annually by 27 million tons
of diatom shell dust transported by transatlantic winds from the
African Sahara, much of it from the Bodélé Depression, which was
once made up of a system of fresh-water lakes.[13][14]
Diatoms
Temporal range:
Light microscopy of a sampling of marine diatoms found living between

crystals of annual sea ice in Antarctica, showing a multiplicity of sizes and
shapes
Scientific classification
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Stramenopiles
Phylum: Gyrista
Subphylum: Ochrophytina
Infraphylum: Diatomista
Superclass: Khakista
Class: Bacillariophyceae
Dangeard, 1933[1]
Synonyms
Diatomea Dumortier, 1821[2]

Diatomophyceae Rabenhorst, 1864[3]
Bacillariae Haeckel, 1878[4]
Bacillariophyta Engler & Gilg, 1919[5]
Diatoms are unicellular organisms: they occur either as solitary cells

or in colonies, which can take the shape of ribbons, fans, zigzags, or
stars. Individual cells range in size from 2 to 200 micrometers.[15] In
the presence of adequate nutrients and sunlight, an assemblage of
living diatoms doubles approximately every 24 hours by asexual
multiple fission; the maximum life span of individual cells is about
six days.[16] Diatoms have two distinct shapes: a few (centric
diatoms) are radially symmetric, while most (pennate diatoms) are
broadly bilaterally symmetric. A unique feature of diatom anatomy is
that they are surrounded by a cell wall made of silica (hydrated
silicon dioxide), called a frustule.[17] These frustules have structural
coloration due to their photonic nanostructure, prompting them to
be described as "jewels of the sea" and "living opals". Movement in
diatoms primarily occurs passively as a result of both ocean
currents and wind-induced water turbulence; however, male
gametes of centric diatoms have flagella, permitting active
movement to seek female gametes. Similar to plants, diatoms
convert light energy to chemical energy by photosynthesis, but their
chloroplasts were acquired in different ways.[18]
Unusually for autotrophic organisms, diatoms possess a urea cycle,
a feature that they share with animals, although this cycle is used to
different metabolic ends in diatoms. The family Rhopalodiaceae also
possess a cyanobacterial endosymbiont called a spheroid body.
This endosymbiont has lost its photosynthetic properties, but has
kept its ability to perform nitrogen fixation, allowing the diatom to fix
atmospheric nitrogen.[19] Other diatoms in symbiosis with nitrogen-
fixing cyanobacteria are among the genera Hemiaulus, Rhizosolenia
and Chaetoceros.[20]
Dinotoms are diatoms that have become endosymbionts inside

dinoflagellates. Research on the dinoflagellates Durinskia baltica and
Glenodinium foliaceum have shown that the endosymbiont event
happened so recently, evolutionarily speaking, that their organelles
and genome are still intact with minimum to no gene loss. The main
difference between these and free living diatoms is that they have
lost their cell wall of silica, making them the only known shell-less
diatoms.[21]
The study of diatoms is a branch of phycology. Diatoms are

classified as eukaryotes, organisms with a nuclear envelope-bound
cell nucleus, that separates them from the prokaryotes archaea and
bacteria. Diatoms are a type of plankton called phytoplankton, the
most common of the plankton types. Diatoms also grow attached to
benthic substrates, floating debris, and on macrophytes. They
comprise an integral component of the periphyton community.[22]
Another classification divides plankton into eight types based on
size: in this scheme, diatoms are classed as microalgae. Several
systems for classifying the individual diatom species exist.
Fossil evidence suggests that diatoms originated during or before

the early Jurassic period, which was about 150 to 200 million years
ago. The oldest fossil evidence for diatoms is a specimen of extant
genus Hemiaulus in Late Jurassic aged amber from Thailand.[23]
Diatoms are used to monitor past and present environmental

conditions, and are commonly used in studies of water quality.
Diatomaceous earth (diatomite) is a collection of diatom shells
found in the earth's crust. They are soft, silica-containing
sedimentary rocks which are easily crumbled into a fine powder and
typically have a particle size of 10 to 200 μm. Diatomaceous earth is
used for a variety of purposes including for water filtration, as a mild
abrasive, in cat litter, and as a dynamite stabilizer.
CC
1:55
Dwindling diatoms and the mixed layer

Earth's oceans teem with microscopic plants called phytoplankton.
But according to a 2015 NASA study, populations of diatoms, the
largest type of phytoplankton algae, have declined more than 1
percent per year from 1998 to 2012. Phytoplankton are an essential
base of the marine food web and absorb carbon dioxide dissolved in
the ocean that originally came from the atmosphere. The tiny
organisms occupy the uppermost layer of ocean water, called the
mixed layer, where waves and currents continually churn, drawing up
nutrients from a deeper layer of water below. Scientists say the
phytoplankton declines observed over the 15-year study period are
due to the mixed layer becoming shallower, which results in fewer
nutrients reaching the diatoms. The reduction in population may
reduce the amount of carbon dioxide drawn out of the atmosphere
and transferred to the deep ocean for long-term storage.[24][25]
0:20
3D-animation of the diatom Corethron sp.

Displays overlays from four fluorescent channels
(a) Green: [DiOC6(3) fluorescence] - stains cellular membranes
indicating the core cell bodies
(b) Cyan: [PLL-A546 fluorescence] - generic counterstain for
visualising eukaryotic cell surfaces
(c) Blue: [Hoechst fluorescence] - stains DNA, identifies nuclei
(d) Red: [chlorophyll autofluorescence] - resolves chloroplasts [26]
The animation starts by overlaying all available fluorescent channels,
and then clarifies the visualisation by switching channels on and off
Overview
Diatoms are protists that form massive annual spring and fall
blooms in aquatic environments and are estimated to be
responsible for about half of photosynthesis in the global oceans.[27]
This predictable annual bloom dynamic fuels higher trophic levels
and initiates delivery of carbon into the deep ocean biome. Diatoms
have complex life history strategies that are presumed to have
contributed to their rapid genetic diversification into ~200,000
species [28] that are distributed between the two major diatom
groups: centrics and pennates.[29][30]
Morphology
Diatoms are generally 2 to 200 micrometers in size,[15] with a few
larger species. Their yellowish-brown chloroplasts, the site of
photosynthesis, are typical of heterokonts, having four cell
membranes and containing pigments such as the carotenoid
fucoxanthin. Individuals usually lack flagella, but they are present in
male gametes of the centric diatoms and have the usual heterokont
structure, including the hairs (mastigonemes) characteristic in other
groups.
Diatoms are often referred as "jewels of the sea" or "living opals" due
to their optical properties.[31] The biological function of this
structural coloration is not clear, but it is speculated that it may be
related to communication, camouflage, thermal exchange and/or UV
protection.[32]
Diatoms build intricate hard but porous cell walls called frustules
composed primarily of silica.[33]: 25–30 This siliceous wall[34] can be
highly patterned with a variety of pores, ribs, minute spines, marginal
ridges and elevations; all of which can be used to delineate genera
and species.
The cell itself consists of two halves, each containing an essentially

flat plate, or valve, and marginal connecting, or girdle band. One half,
the hypotheca, is slightly smaller than the other half, the epitheca.
Diatom morphology varies. Although the shape of the cell is typically
circular, some cells may be triangular, square, or elliptical. Their
distinguishing feature is a hard mineral shell or frustule composed
of opal (hydrated, polymerized silicic acid).
Representation of a diatom
1. Nucleus; holds the genetic material
2. Nucleolus; location of the chromosomes
3. Golgi apparatus; modifies proteins and sends them out of

the cell
4. Cell wall; outer membrane of the cell
5. Pyrenoid; center of carbon fixation
6. Chromatophore; pigment carrying membrane structure
7. Vacuoles; vesicle of a cell that contains fluid bound by a

membrane
8. Cytoplasmic strands; hold the nucleus
9. Mitochondria; create ATP (energy) for the cell
10. Valves/Striae; allow nutrients in, and waste out, of the cell
Intricate structures of the diatom
a. Areolae (hexagonal or polygonal boxlike perforation with a sieve
present on the surface of diatom)
b. Striae (pores, punctae, spots or dots in a line on the surface)
c. Raphe (slit in the valves)
d. Central nodule (thickening of wall at the midpoint of raphe)
e. Stigmata (holes through valve surface which looks rounded

externally but with a slit like internal)
f. Punctae (spots or small perforations on the surface)
g. Polar nodules (thickening of wall at the distal ends of the

raphe)[35][36]
Selections from Ernst Haeckel's 1904

Kunstformen der Natur (Art Forms of Nature),
showing pennate (left) and centric (right)
frustules.
Diatoms are divided into two groups that are distinguished by the
shape of the frustule: the centric diatoms and the pennate diatoms.
Pennate diatoms are bilaterally symmetric. Each one of their valves

have openings that are slits along the raphes and their shells are
typically elongated parallel to these raphes. They generate cell
movement through cytoplasm that streams along the raphes,
always moving along solid surfaces.
Centric diatoms are radially symmetric. They are composed of upper
and lower valves – epitheca and hypotheca – each consisting of a
valve and a girdle band that can easily slide underneath each other
and expand to increase cell content over the diatoms progression.
The cytoplasm of the centric diatom is located along the inner
surface of the shell and provides a hollow lining around the large
vacuole located in the center of the cell. This large, central vacuole
is filled by a fluid known as "cell sap" which is similar to seawater
but varies with specific ion content. The cytoplasmic layer is home
to several organelles, like the chloroplasts and mitochondria. Before
the centric diatom begins to expand, its nucleus is at the center of
one of the valves and begins to move towards the center of the
cytoplasmic layer before division is complete. Centric diatoms have
a variety of shapes and sizes, depending on from which axis the
shell extends, and if spines are present.
Shape classification of diatom frustules. The

images are 3D models. The actual sizes of the
frustules are about 10–80 μm.[37]
Structure of a centric diatom frustule [37]

Silicification
Diatom cells are contained within a unique silica cell wall known as
a frustule made up of two valves called thecae, that typically overlap
one another.[38] The biogenic silica composing the cell wall is
synthesised intracellularly by the polymerisation of silicic acid
monomers. This material is then extruded to the cell exterior and
added to the wall. In most species, when a diatom divides to
produce two daughter cells, each cell keeps one of the two-halves
and grows a smaller half within it. As a result, after each division
cycle, the average size of diatom cells in the population gets
smaller. Once such cells reach a certain minimum size, rather than
simply divide, they reverse this decline by forming an auxospore.
This expands in size to give rise to a much larger cell, which then
returns to size-diminishing divisions. Auxospore production is
almost always linked to meiosis and sexual reproduction.
Pennate diatom from an Arctic meltpond, infected

with two chytrid-like [zoo-]sporangium fungal
pathogens (in false-colour red). Scale bar = 10 μm.[39]
Light microscopy of a living diatom. Numbered
graduations are 10 micrometres apart
The exact mechanism of transferring silica absorbed by the diatom

to the cell wall is unknown. Much of the sequencing of diatom genes
comes from the search for the mechanism of silica uptake and
deposition in nano-scale patterns in the frustule. The most success
in this area has come from two species, Thalassiosira pseudonana,
which has become the model species, as the whole genome was
sequenced and methods for genetic control were established, and
Cylindrotheca fusiformis, in which the important silica deposition
proteins silaffins were first discovered.[40] Silaffins, sets of
polycationic peptides, were found in C. fusiformis cell walls and can
generate intricate silica structures. These structures demonstrated
pores of sizes characteristic to diatom patterns. When T.
pseudonana underwent genome analysis it was found that it
encoded a urea cycle, including a higher number of polyamines than
most genomes, as well as three distinct silica transport genes.[41] In
a phylogenetic study on silica transport genes from 8 diverse groups
of diatoms, silica transport was found to generally group with
species.[40] This study also found structural differences between the
silica transporters of pennate (bilateral symmetry) and centric
(radial symmetry) diatoms. The sequences compared in this study
were used to create a diverse background in order to identify
residues that differentiate function in the silica deposition process.
Additionally, the same study found that a number of the regions
were conserved within species, likely the base structure of silica
transport.
These silica transport proteins are unique to diatoms, with no

homologs found in other species, such as sponges or rice. The
divergence of these silica transport genes is also indicative of the
structure of the protein evolving from two repeated units composed
of five membrane bound segments, which indicates either gene
duplication or dimerization.[40] The silica deposition that takes place
from the membrane bound vesicle in diatoms has been
hypothesized to be a result of the activity of silaffins and long chain
polyamines. This Silica Deposition Vesicle (SDV) has been
characterized as an acidic compartment fused with Golgi-derived
vesicles.[42] These two protein structures have been shown to create
sheets of patterned silica in-vivo with irregular pores on the scale of
diatom frustules. One hypothesis as to how these proteins work to
create complex structure is that residues are conserved within the
SDV's, which is unfortunately difficult to identify or observe due to
the limited number of diverse sequences available. Though the
exact mechanism of the highly uniform deposition of silica is as yet
unknown, the Thalassiosira pseudonana genes linked to silaffins are
being looked to as targets for genetic control of nanoscale silica
deposition.
The ability of diatoms to make silica-based cell walls has been the
subject of fascination for centuries. It started with a microscopic
observation by an anonymous English country nobleman in 1703,
who observed an object that looked like a chain of regular
parallelograms and debated whether it was just crystals of salt, or a
plant.[43] The viewer decided that it was a plant because the
parallelograms didn't separate upon agitation, nor did they vary in
appearance when dried or subjected to warm water (in an attempt to
dissolve the "salt"). Unknowingly, the viewer's confusion captured
the essence of diatoms—mineral utilizing plants. It is not clear when
it was determined that diatom cell walls are made of silica, but in
1939 a seminal reference characterized the material as silicic acid in
a "subcolloidal" state[44] Identification of the main chemical
component of the cell wall spurred investigations into how it was
made. These investigations have involved, and been propelled by,
diverse approaches including, microscopy, chemistry, biochemistry,
material characterisation, molecular biology, 'omics, and transgenic
approaches. The results from this work have given a better
understanding of cell wall formation processes, establishing
fundamental knowledge which can be used to create models that
contextualise current findings and clarify how the process works.[45]
The process of building a mineral-based cell wall inside the cell, then
exporting it outside, is a massive event that must involve large
numbers of genes and their protein products. The act of building
and exocytosing this large structural object in a short time period,
synched with cell cycle progression, necessitates substantial
physical movements within the cell as well as dedication of a
significant proportion of the cell's biosynthetic capacities.[45]
The first characterisations of the biochemical processes and

components involved in diatom silicification were made in the late
1990s.[46][47][48] These were followed by insights into how higher
order assembly of silica structures might occur.[49][50][51] More
recent reports describe the identification of novel components
involved in higher order processes, the dynamics documented
through real-time imaging, and the genetic manipulation of silica
structure.[52][53] The approaches established in these recent works
provide practical avenues to not only identify the components
involved in silica cell wall formation but to elucidate their
interactions and spatio-temporal dynamics. This type of holistic
understanding will be necessary to achieve a more complete
understanding of cell wall synthesis.[45]
Behaviour
Chaetoceros willei Chaetoceros furcillatus J.W.Bailey,

Gran, 1897 1856
Most centric and araphid pennate diatoms are nonmotile, and their
relatively dense cell walls cause them to readily sink. Planktonic
forms in open water usually rely on turbulent mixing of the upper
layers of the oceanic waters by the wind to keep them suspended in
sunlit surface waters. Many planktonic diatoms have also evolved
features that slow their sinking rate, such as spines or the ability to
grow in colonial chains.[54] These adaptations increase their surface
area to volume ratio and drag, allowing them to stay suspended in
the water column longer. Individual cells may regulate buoyancy via
an ionic pump.[55]
Some pennate diatoms are capable of a type of locomotion called

"gliding", which allows them to move across surfaces via adhesive
mucilage secreted through a seamlike structure called the
raphe.[56][57] In order for a diatom cell to glide, it must have a solid
substrate for the mucilage to adhere to.
Cells are solitary or united into colonies of various kinds, which may
be linked by siliceous structures; mucilage pads, stalks or tubes;
amorphous masses of mucilage; or by threads of chitin
(polysaccharide), which are secreted through strutted processes of
the cell.
Planktonic diatoms such as Thalassiosira sp. (56-62),

Asteromphalus sp. (63), Aulacoseira sp. (64-66), and
Chaetoceros (see twin image above) often grow in
chains, and have features such as spines which slow
sinking rates by increasing drag.
Some Thalassiosira diatoms form chain-like colonies, like these

collected near the Antarctic peninsula coast by the schooner of the
Tara Oceans Expedition for plankton research.
This projection of a stack of confocal images shows the diatoms' cell
wall (cyan), chloroplasts (red), DNA (blue), membranes and
organelles (green).
Life cycle
Sexual reproduction
Centric diatom (oogamy) Pennate diatom (morphological

isogamy, physiological anisogamy)
Reproduction and cell size
Reproduction among these organisms is asexual by binary fission,

during which the diatom divides into two parts, producing two "new"
diatoms with identical genes. Each new organism receives one of
the two frustules – one larger, the other smaller – possessed by the
parent, which is now called the epitheca; and is used to construct a
second, smaller frustule, the hypotheca. The diatom that received
the larger frustule becomes the same size as its parent, but the
diatom that received the smaller frustule remains smaller than its
parent. This causes the average cell size of this diatom population
to decrease.[15] It has been observed, however, that certain taxa
have the ability to divide without causing a reduction in cell size.[58]
Nonetheless, in order to restore the cell size of a diatom population
for those that do endure size reduction, sexual reproduction and
auxospore formation must occur.[15]
Cell division
Vegetative cells of diatoms are diploid (2N) and so meiosis can take
place, producing male and female gametes which then fuse to form
the zygote. The zygote sheds its silica theca and grows into a large
sphere covered by an organic membrane, the auxospore. A new
diatom cell of maximum size, the initial cell, forms within the
auxospore thus beginning a new generation. Resting spores may
also be formed as a response to unfavourable environmental
conditions with germination occurring when conditions improve.[33]
A defining characteristic of all diatoms is their restrictive and

bipartite silica cell wall that causes them to progressively shrink
during asexual cell division. At a critically small cell size and under
certain conditions, auxosporulation restitutes cell size and prevents
clonal death.[59][60][61][62][63] The entire lifecycles of only a few
diatoms have been described and rarely have sexual events been
captured in the environment.[30]
Sperm motility
Diatoms are mostly non-motile; however, sperm found in some

species can be flagellated, though motility is usually limited to a
gliding motion.[33] In centric diatoms, the small male gametes have
one flagellum while the female gametes are large and non-motile
(oogamous). Conversely, in pennate diatoms both gametes lack
flagella (isogamous).[15] Certain araphid species, that is pennate
diatoms without a raphe (seam), have been documented as
anisogamous and are, therefore, considered to represent a
transitional stage between centric and raphid pennate diatoms,
diatoms with a raphe.[58]
Degradation by microbes
Certain species of bacteria in oceans and lakes can accelerate the

rate of dissolution of silica in dead and living diatoms by using
hydrolytic enzymes to break down the organic algal material.[64][65]
Ecology
Regions of high abundance of diatoms in the ocean
Diatom dominance (as a percentage of total cell counts)

versus silicate concentration [66]
Distribution
Diatoms are a widespread group and can be found in the oceans, in

fresh water, in soils, and on damp surfaces. They are one of the
dominant components of phytoplankton in nutrient-rich coastal
waters and during oceanic spring blooms, since they can divide
more rapidly than other groups of phytoplankton.[67] Most live
pelagically in open water, although some live as surface films at the
water-sediment interface (benthic), or even under damp
atmospheric conditions. They are especially important in oceans,
where they contribute an estimated 45% of the total oceanic primary
production of organic material.[68] Spatial distribution of marine
phytoplankton species is restricted both horizontally and
vertically.[69][33]
Growth
Planktonic diatoms in freshwater and marine environments typically

exhibit a "boom and bust" (or "bloom and bust") lifestyle. When
conditions in the upper mixed layer (nutrients and light) are
favourable (as at the spring), their competitive edge and rapid
growth rate[67] enables them to dominate phytoplankton
communities ("boom" or "bloom"). As such they are often classed as
opportunistic r-strategists (i.e. those organisms whose ecology is
defined by a high growth rate, r).
Impact
The freshwater diatom Didymosphenia geminata, commonly known

as Didymo, causes severe environmental degradation in water-
courses where it blooms, producing large quantities of a brown jelly-
like material called "brown snot" or "rock snot". This diatom is native
to Europe and is an invasive species both in the antipodes and in
parts of North America.[70][71] The problem is most frequently
recorded from Australia and New Zealand.[72]
When conditions turn unfavourable, usually upon depletion of
nutrients, diatom cells typically increase in sinking rate and exit the
upper mixed layer ("bust"). This sinking is induced by either a loss of
buoyancy control, the synthesis of mucilage that sticks diatoms
cells together, or the production of heavy resting spores. Sinking out
of the upper mixed layer removes diatoms from conditions
unfavourable to growth, including grazer populations and higher
temperatures (which would otherwise increase cell metabolism).
Cells reaching deeper water or the shallow seafloor can then rest
until conditions become more favourable again. In the open ocean,
many sinking cells are lost to the deep, but refuge populations can
persist near the thermocline.
Ultimately, diatom cells in these resting populations re-enter the

upper mixed layer when vertical mixing entrains them. In most
circumstances, this mixing also replenishes nutrients in the upper
mixed layer, setting the scene for the next round of diatom blooms.
In the open ocean (away from areas of continuous upwelling[73]),
this cycle of bloom, bust, then return to pre-bloom conditions
typically occurs over an annual cycle, with diatoms only being
prevalent during the spring and early summer. In some locations,
however, an autumn bloom may occur, caused by the breakdown of
summer stratification and the entrainment of nutrients while light
levels are still sufficient for growth. Since vertical mixing is
increasing, and light levels are falling as winter approaches, these
blooms are smaller and shorter-lived than their spring equivalents.
In the open ocean, the diatom (spring) bloom is typically ended by a

shortage of silicon. Unlike other minerals, the requirement for silicon
is unique to diatoms and it is not regenerated in the plankton
ecosystem as efficiently as, for instance, nitrogen or phosphorus
nutrients. This can be seen in maps of surface nutrient
concentrations – as nutrients decline along gradients, silicon is
usually the first to be exhausted (followed normally by nitrogen then
phosphorus).
Because of this bloom-and-bust cycle, diatoms are believed to play a

disproportionately important role in the export of carbon from
oceanic surface waters[73][74] (see also the biological pump).
Significantly, they also play a key role in the regulation of the
biogeochemical cycle of silicon in the modern ocean.[68][75]
Reason for success
Diatoms are ecologically successful, and occur in virtually every

environment that contains water – not only oceans, seas, lakes, and
streams, but also soil and wetlands. The use of silicon by diatoms is
believed by many researchers to be the key to this ecological
success. Raven (1983)[76] noted that, relative to organic cell walls,
silica frustules require less energy to synthesize (approximately 8%
of a comparable organic wall), potentially a significant saving on the
overall cell energy budget. In a now classic study, Egge and Aksnes
(1992)[66] found that diatom dominance of mesocosm communities
was directly related to the availability of silicic acid – when
concentrations were greater than 2 μmol m−3, they found that
diatoms typically represented more than 70% of the phytoplankton
community. Other researchers[77] have suggested that the biogenic
silica in diatom cell walls acts as an effective pH buffering agent,
facilitating the conversion of bicarbonate to dissolved CO2 (which is
more readily assimilated). More generally, notwithstanding these
possible advantages conferred by their use of silicon, diatoms
typically have higher growth rates than other algae of the same
corresponding size.[67]
Sources for collection
Diatoms can be obtained from multiple sources.[78] Marine diatoms

can be collected by direct water sampling, and benthic forms can be
secured by scraping barnacles, oyster and other shells. Diatoms are
frequently present as a brown, slippery coating on submerged
stones and sticks, and may be seen to "stream" with river current.
The surface mud of a pond, ditch, or lagoon will almost always yield
some diatoms. Living diatoms are often found clinging in great
numbers to filamentous algae, or forming gelatinous masses on
various submerged plants. Cladophora is frequently covered with
Cocconeis, an elliptically shaped diatom; Vaucheria is often covered
with small forms. Since diatoms form an important part of the food
of molluscs, tunicates, and fishes, the alimentary tracts of these
animals often yield forms that are not easily secured in other ways.
Diatoms can be made to emerge by filling a jar with water and mud,
wrapping it in black paper and letting direct sunlight fall on the
surface of the water. Within a day, the diatoms will come to the top
in a scum and can be isolated.[78]
Biogeochemistry
S
il
i
c
a
c
y
c
l
e
The modern oceanic silicon cycle
Fluxes are in T mol Si y−1 (28 million metric tons of silicon per year)
The diagram shows the major fluxes of silicon in the current ocean.
Most biogenic silica in the ocean (silica produced by biological
activity) comes from diatoms. Diatoms extract dissolved silicic acid
from surface waters as they grow, and return it to the water column
when they die. Inputs of silicon arrive from above via aeolian dust,
from the coasts via rivers, and from below via seafloor sediment
recycling, weathering, and hydrothermal activity.[75]
Although diatoms may have existed since the Triassic, the timing of
their ascendancy and "take-over" of the silicon cycle occurred more
recently. Prior to the Phanerozoic (before 544 Ma), it is believed that
microbial or inorganic processes weakly regulated the ocean's
silicon cycle.[79][80][81] Subsequently, the cycle appears dominated
(and more strongly regulated) by the radiolarians and siliceous
sponges, the former as zooplankton, the latter as sedentary filter-
feeders primarily on the continental shelves.[82] Within the last 100
My, it is thought that the silicon cycle has come under even tighter
control, and that this derives from the ecological ascendancy of the
diatoms.
However, the precise timing of the "take-over" remains unclear, and

different authors have conflicting interpretations of the fossil record.
Some evidence, such as the displacement of siliceous sponges
from the shelves,[83] suggests that this takeover began in the
Cretaceous (146 Ma to 66 Ma), while evidence from radiolarians
suggests "take-over" did not begin until the Cenozoic (66 Ma to
present).[84]
C
a
r
b
o
n
c
y
c
l
e Ocean carbon cycle and diatom carbon dioxide concentration
mechanisms [85]
The diagram depicts some mechanisms by which marine diatoms

contribute to the biological carbon pump and influence the ocean
carbon cycle. The anthropogenic CO2 emission to the atmosphere
(mainly generated by fossil fuel burning and deforestation) is nearly
11 gigatonne carbon (GtC) per year, of which almost 2.5 GtC is taken
up by the surface ocean. In surface seawater (pH 8.1–8.4),
bicarbonate (HCO−3) and carbonate ions (CO2−
3 ) constitute nearly 90
and <10% of dissolved inorganic carbon (DIC) respectively, while
dissolved CO2 (CO2 aqueous) contributes <1%. Despite this low level
of CO2 in the ocean and its slow diffusion rate in water, diatoms fix
10–20 GtC annually via photosynthesis thanks to their carbon
dioxide concentrating mechanisms, allowing them to sustain marine
food chains. In addition, 0.1–1% of this organic material produced in
the euphotic layer sinks down as particles, thus transferring the
surface carbon toward the deep ocean and sequestering
atmospheric CO2 for thousands of years or longer. The remaining
organic matter is remineralized through respiration. Thus, diatoms
are one of the main players in this biological carbon pump, which is
arguably the most important biological mechanism in the Earth
System allowing CO2 to be removed from the carbon cycle for very
long period.[86][85]
U
r
e
a
c
y
c
l
eMitochondrial urea cycle in a generic diatom cell and the potential fates
of urea cycle intermediates [87]
A feature of diatoms is the urea cycle, which links them
evolutionarily to animals. In 2011, Allen et al. established that
diatoms have a functioning urea cycle. This result was significant,
since prior to this, the urea cycle was thought to have originated
with the metazoans which appeared several hundreds of millions of
years before the diatoms. Their study demonstrated that while
diatoms and animals use the urea cycle for different ends, they are
seen to be evolutionarily linked in such a way that animals and
plants are not.[88]
While often overlooked in photosynthetic organisms, the

mitochondria also play critical roles in energy balance. Two
nitrogen-related pathways are relevant and they may also change
under ammonium (NH+4) nutrition compared with nitrate (NO−3)
nutrition. First, in diatoms, and likely some other algae, there is a
urea cycle.[89][90][91] The long-known function of the urea cycle in
animals is to excrete excess nitrogen produced by amino acid
catabolism; like photorespiration, the urea cycle had long been
considered a waste pathway. However, in diatoms the urea cycle
appears to play a role in exchange of nutrients between the
mitochondria and the cytoplasm, and potentially the plastid [92] and
may help to regulate ammonium metabolism.[89][90] Because of this
cycle, marine diatoms, in contrast to chlorophytes, also have
acquired a mitochondrial urea transporter and, in fact, based on
bioinformatics, a complete mitochondrial GS-GOGAT cycle has been
hypothesised.[90][87]
Other
Diatoms are mainly photosynthetic; however a few are obligate

heterotrophs and can live in the absence of light provided an
appropriate organic carbon source is available.[93][94]
Photosynthetic diatoms that find themselves in an environment

absent of oxygen and/or sunlight can switch to anaerobic
respiration known as nitrate respiration (DNRA), and stay dormant
for up till months and decades.[95][96]
Major pigments of diatoms are chlorophylls a and c, beta-carotene,

fucoxanthin, diatoxanthin and diadinoxanthin.[15]
Taxonomy
Light microscopy of several species

of living freshwater diatoms
Centric diatom
Linked diatoms
Thalassiosirales
Stephanodiscus hantzschii
Coscinodiscophyceae
Isthmia nervosaIsthmia nervosa
Coscinodiscophyceae
Odontella aurita
Diatoms belong to a large group of protists, many of which contain

plastids rich in chlorophylls a and c. The group has been variously
referred to as heterokonts, chrysophytes, chromists or
stramenopiles. Many are autotrophs such as golden algae and kelp;
and heterotrophs such as water moulds, opalinids, and actinophryid
heliozoa. The classification of this area of protists is still unsettled.
In terms of rank, they have been treated as a division, phylum,
kingdom, or something intermediate to those. Consequently,
diatoms are ranked anywhere from a class, usually called
Diatomophyceae or Bacillariophyceae, to a division (=phylum),
usually called Bacillariophyta, with corresponding changes in the
ranks of their subgroups.
Genera and species
An estimated 20,000 extant diatom species are believed to exist, of

which around 12,000 have been named to date according to Guiry,
2012[97] (other sources give a wider range of estimates[15][98][99][100]).
Around 1,000–1,300 diatom genera have been described, both
extant and fossil,[101][102] of which some 250–300 exist only as
fossils.[103]
Classes and orders
For many years the diatoms—treated either as a class

(Bacillariophyceae) or a phylum (Bacillariophyta)—were divided into
just 2 orders, corresponding to the centric and the pennate diatoms
(Centrales and Pennales). This classification was extensively
overhauled by Round, Crawford and Mann in 1990 who treated the
diatoms at a higher rank (division, corresponding to phylum in
zoological classification), and promoted the major classification
units to classes, maintaining the centric diatoms as a single class
Coscinodiscophyceae, but splitting the former pennate diatoms into
2 separate classes, Fragilariophyceae and Bacillariophyceae (the
latter older name retained but with an emended definition), between
them encompassing 45 orders, the majority of them new.
Today (writing at mid 2020) it is recognised that the 1990 system of

Round et al. is in need of revision with the advent of newer
molecular work, however the best system to replace it is unclear,
and current systems in widespread use such as AlgaeBase, the
World Register of Marine Species and its contributing database
DiatomBase, and the system for "all life" represented in Ruggiero et
al., 2015, all retain the Round et al. treatment as their basis, albeit
with diatoms as a whole treated as a class rather than
division/phylum, and Round et al.'s classes reduced to subclasses,
for better agreement with the treatment of phylogenetically adjacent
groups and their containing taxa. (For references refer the individual
sections below).
One proposal, by Linda Medlin and co-workers commencing in 2004,
is for some of the centric diatom orders considered more closely
related to the pennates to be split off as a new class, Mediophyceae,
itself more closely aligned with the pennate diatoms than the
remaining centrics. This hypothesis—later designated the
Coscinodiscophyceae-Mediophyceae-Bacillariophyceae, or
Coscinodiscophyceae+(Mediophyceae+Bacillariophyceae) (CMB)
hypothesis—has been accepted by D.G. Mann among others, who
uses it as the basis for the classification of diatoms as presented in
Adl. et al.'s series of syntheses (2005, 2012, 2019), and also in the
Bacillariophyta chapter of the 2017 Handbook of the Protists edited
by Archibald et al., with some modifications reflecting the apparent
non-monophyly of Medlin et al. original "Coscinodiscophyceae".
Meanwhile, a group led by E.C. Theriot favours a different
hypothesis of phylogeny, which has been termed the structural
gradation hypothesis (SGH) and does not recognise the
Mediophyceae as a monophyletic group, while another analysis, that
of Parks et al., 2018, finds that the radial centric diatoms (Medlin et
al.'s Coscinodiscophyceae) are not monophyletic, but supports the
monophyly of Mediophyceae minus Attheya, which is an anomalous
genus. Discussion of the relative merits of these conflicting
schemes continues by the various parties involved.[104][105][106][107]
Adl et al., 2019 treatment
In 2019, Adl et al.[108] presented the following classification of

diatoms, while noting: "This revision reflects numerous advances in
the phylogeny of the diatoms over the last decade. Due to our poor
taxon sampling outside of the Mediophyceae and pennate diatoms,
and the known and anticipated diversity of all diatoms, many clades
appear at a high classification level (and the higher level
classification is rather flat)." This classification treats diatoms as a
phylum (Diatomeae/Bacillariophyta), accepts the class
Mediophyceae of Medlin and co-workers, introduces new subphyla
and classes for a number of otherwise isolated genera, and re-ranks
a number of previously established taxa as subclasses, but does not
list orders or families. Inferred ranks have been added for clarity
(Adl. et al. do not use ranks, but the intended ones in this portion of
the classification are apparent from the choice of endings used,
within the system of botanical nomenclature employed).
Clade Diatomista Derelle et al. 2016, emend. Cavalier-Smith 2017

(diatoms plus a subset of other ochrophyte groups)
Phylum Diatomeae Dumortier 1821 [= Bacillariophyta Haeckel
1878] (diatoms)
Subphylum Leptocylindrophytina D.G. Mann in Adl et al.
2019
Class Leptocylindrophyceae D.G. Mann in Adl et al.
2019 (Leptocylindrus, Tenuicylindrus)
Class Corethrophyceae D.G. Mann in Adl et al. 2019
(Corethron)
Subphylum Ellerbeckiophytina D.G. Mann in Adl et al.
2019 (Ellerbeckia)
Subphylum Probosciophytina D.G. Mann in Adl et al.
2019 (Proboscia)
Subphylum Melosirophytina D.G. Mann in Adl et al. 2019
(Aulacoseira, Melosira, Hyalodiscus, Stephanopyxis,
Paralia, Endictya)
Subphylum Coscinodiscophytina Medlin & Kaczmarska
2004, emend. (Actinoptychus, Coscinodiscus,
Actinocyclus, Asteromphalus, Aulacodiscus, Stellarima)
Subphylum Rhizosoleniophytina D.G. Mann in Adl et al.
2019 (Guinardia, Rhizosolenia, Pseudosolenia)
Subphylum Arachnoidiscophytina D.G. Mann in Adl et al.
2019 (Arachnoidiscus)
Subphylum Bacillariophytina Medlin & Kaczmarska 2004,
emend.
Class Mediophyceae Jouse & Proshkina-Lavrenko in
Medlin & Kaczmarska 2004
Subclass Chaetocerotophycidae Round & R.M.
Crawford in Round et al. 1990, emend.
Subclass Lithodesmiophycidae Round & R.M.
Subclass Thalassiosirophycidae Round & R.M.
Crawford in Round et al. 1990
Subclass Cymatosirophycidae Round & R.M.
Crawford in Round et al. 1990
Subclass Odontellophycidae D.G. Mann in Adl et
al. 2019
Subclass Chrysanthemodiscophycidae D.G.
Mann in Adl et al. 2019
Class Biddulphiophyceae D.G. Mann in Adl et al.
2019
Subclass Biddulphiophycidae Round and R.M.
Biddulphiophyceae incertae sedis (Attheya)
Class Bacillariophyceae Haeckel 1878, emend.
Bacillariophyceae incertae sedis (Striatellaceae)
Subclass Urneidophycidae Medlin 2016
Subclass Fragilariophycidae Round in Round,
Crawford & Mann 1990, emend.
Subclass Bacillariophycidae D.G. Mann in
Round, Crawford & Mann 1990, emend.
See taxonomy of diatoms for more details.
Gallery
Scanning electron microscope images
Diatom Surirella spiralis
Diatoms Thalassiosira sp. on a membrane filter, pore size 0.4 μm.

Diatom Paralia sulcata.
Diatom Achanthes trinodis
External video
Bacillaria: Distractingly Beautiful Crystal Colonies (https://www.youtube.
com/watch?v=Nysdq_plUTg&ab_channel=JourneytotheMicrocosmos) –
Journey to the Microcosmos:
Stand-alone cell of Bacillaria paxillifer
Colonial group of Bacillaria paxillifer
Three diatom species were sent to the International Space Station,

including the huge (6 mm length) diatoms of Antarctica and the
exclusive colonial diatom, Bacillaria paradoxa. The cells of Bacillaria
moved next to each other in partial but opposite synchrony by a
microfluidics method.[109]
Evolution and fossil record
Origin
Heterokont chloroplasts appear to derive from those of red algae,

rather than directly from prokaryotes as occurred in plants. This
suggests they had a more recent origin than many other algae.
However, fossil evidence is scant, and only with the evolution of the
diatoms themselves do the heterokonts make a serious impression
on the fossil record.
Earliest fossils
The earliest known fossil diatoms date from the early Jurassic
(~185 Ma ago),[110] although the molecular clock[110] and
sedimentary[111] evidence suggests an earlier origin. It has been
suggested that their origin may be related to the end-Permian mass
extinction (~250 Ma), after which many marine niches were
opened.[112] The gap between this event and the time that fossil
diatoms first appear may indicate a period when diatoms were
unsilicified and their evolution was cryptic.[113] Since the advent of
silicification, diatoms have made a significant impression on the
fossil record, with major fossil deposits found as far back as the
early Cretaceous, and with some rocks such as diatomaceous earth,
being composed almost entirely of them.
Relation to grasslands
The expansion of grassland biomes and the evolutionary radiation

of grasses during the Miocene is believed to have increased the flux
of soluble silicon to the oceans, and it has been argued that this
promoted the diatoms during the Cenozoic era.[114][115] Recent work
suggests that diatom success is decoupled from the evolution of
grasses, although both diatom and grassland diversity increased
strongly from the middle Miocene.[116]
Relation to climate
Diatom diversity over the Cenozoic has been very sensitive to global
temperature, particularly to the equator-pole temperature gradient.
Warmer oceans, particularly warmer polar regions, have in the past
been shown to have had substantially lower diatom diversity. Future
warm oceans with enhanced polar warming, as projected in global-
warming scenarios,[117] could thus in theory result in a significant
loss of diatom diversity, although from current knowledge it is
impossible to say if this would occur rapidly or only over many tens
of thousands of years.[116]
Method of investigation
The fossil record of diatoms has largely been established through

the recovery of their siliceous frustules in marine and non-marine
sediments. Although diatoms have both a marine and non-marine
stratigraphic record, diatom biostratigraphy, which is based on time-
constrained evolutionary originations and extinctions of unique taxa,
is only well developed and widely applicable in marine systems. The
duration of diatom species ranges have been documented through
the study of ocean cores and rock sequences exposed on land.[118]
Where diatom biozones are well established and calibrated to the
geomagnetic polarity time scale (e.g., Southern Ocean, North
Pacific, eastern equatorial Pacific), diatom-based age estimates
may be resolved to within <100,000 years, although typical age
resolution for Cenozoic diatom assemblages is several hundred
thousand years.
Diatoms preserved in lake sediments are widely used for

paleoenvironmental reconstructions of Quaternary climate,
especially for closed-basin lakes which experience fluctuations in
water depth and salinity.
Isotope records
Intricate silicate (glass) shell, 32-40

million years old, of a diatom
microfossil
When diatoms die their shells (frustules) can settle on the seafloor
and become microfossils. Over time, these microfossils become
buried as opal deposits in the marine sediment. Paleoclimatology is
the study of past climates. Proxy data is used in order to relate
elements collected in modern-day sedimentary samples to climatic
and oceanic conditions in the past. Paleoclimate proxies refer to
preserved or fossilized physical markers which serve as substitutes
for direct meteorological or ocean measurements.[119] An example
of proxies is the use of diatom isotope records of δ13C, δ18O, δ30Si
(δ13Cdiatom, δ18Odiatom, and δ30Sidiatom). In 2015, Swann and
Snelling used these isotope records to document historic changes in
the photic zone conditions of the north-west Pacific Ocean,
including nutrient supply and the efficiency of the soft-tissue
biological pump, from the modern day back to marine isotope stage
5e, which coincides with the last interglacial period. Peaks in opal
productivity in the marine isotope stage are associated with the
breakdown of the regional halocline stratification and increased
nutrient supply to the photic zone.[120]
The initial development of the halocline and stratified water column

has been attributed to the onset of major Northern Hemisphere
glaciation at 2.73 Ma, which increased the flux of freshwater to the
region, via increased monsoonal rainfall and/or glacial meltwater,
and sea surface temperatures.[121][122][123][124] The decrease of
abyssal water upwelling associated with this may have contributed
to the establishment of globally cooler conditions and the expansion
of glaciers across the Northern Hemisphere from 2.73 Ma.[122] While
the halocline appears to have prevailed through the late Pliocene
and early Quaternary glacial–interglacial cycles,[125] other studies
have shown that the stratification boundary may have broken down
in the late Quaternary at glacial terminations and during the early
part of interglacials.[126][127][128][129][130][120]
Diversification
The Cretaceous record of diatoms is limited, but recent studies

reveal a progressive diversification of diatom types. The
Cretaceous–Paleogene extinction event, which in the oceans
dramatically affected organisms with calcareous skeletons, appears
to have had relatively little impact on diatom evolution.[131]
Turnover
Although no mass extinctions of marine diatoms have been

observed during the Cenozoic, times of relatively rapid evolutionary
turnover in marine diatom species assemblages occurred near the
Paleocene–Eocene boundary,[132] and at the Eocene–Oligocene
boundary.[133] Further turnover of assemblages took place at various
times between the middle Miocene and late Pliocene,[134] in
response to progressive cooling of polar regions and the
development of more endemic diatom assemblages.
A global trend toward more delicate diatom frustules has been
noted from the Oligocene to the Quaternary.[118] This coincides with
an increasingly more vigorous circulation of the ocean's surface and
deep waters brought about by increasing latitudinal thermal
gradients at the onset of major ice sheet expansion on Antarctica
and progressive cooling through the Neogene and Quaternary
towards a bipolar glaciated world. This caused diatoms to take in
less silica for the formation of their frustules. Increased mixing of
the oceans renews silica and other nutrients necessary for diatom
growth in surface waters, especially in regions of coastal and
oceanic upwelling.
Genetics
Phaeodactylum tricornutum is widely

used as a model organism
Expressed sequence tagging
In 2002, the first insights into the properties of the Phaeodactylum

tricornutum gene repertoire were described using 1,000 expressed
sequence tags (ESTs).[135] Subsequently, the number of ESTs was
extended to 12,000 and the diatom EST database was constructed
for functional analyses.[136] These sequences have been used to
make a comparative analysis between P. tricornutum and the
putative complete proteomes from the green alga Chlamydomonas
reinhardtii, the red alga Cyanidioschyzon merolae, and the diatom
Thalassiosira pseudonana.[137] The diatom EST database now
consists of over 200,000 ESTs from P. tricornutum (16 libraries) and
T. pseudonana (7 libraries) cells grown in a range of different
conditions, many of which correspond to different abiotic
stresses.[138]
Genome sequencing
Thalassiosira pseudonana was the

first eukaryotic marine
phytoplankton to have its genome
sequenced
In 2004, the entire genome of the centric diatom, Thalassiosira

pseudonana (32.4 Mb) was sequenced,[139] followed in 2008 with the
sequencing of the pennate diatom, Phaeodactylum tricornutum (27.4
Mb).[140] Comparisons of the two reveal that the P. tricornutum
genome includes fewer genes (10,402 opposed to 11,776) than T.
pseudonana; no major synteny (gene order) could be detected
between the two genomes. T. pseudonana genes show an average
of ~1.52 introns per gene as opposed to 0.79 in P. tricornutum,
suggesting recent widespread intron gain in the centric
diatom.[140][141] Despite relatively recent evolutionary divergence (90
million years), the extent of molecular divergence between centrics
and pennates indicates rapid evolutionary rates within the
Bacillariophyceae compared to other eukaryotic groups.[140]
Comparative genomics also established that a specific class of
transposable elements, the Diatom Copia-like retrotransposons (or
CoDis), has been significantly amplified in the P. tricornutum genome
with respect to T. pseudonana, constituting 5.8 and 1% of the
respective genomes.[142]
Endosymbiotic gene transfer
Diatom genomics brought much information about the extent and

dynamics of the endosymbiotic gene transfer (EGT) process.
Comparison of the T. pseudonana proteins with homologs in other
organisms suggested that hundreds have their closest homologs in
the Plantae lineage. EGT towards diatom genomes can be illustrated
by the fact that the T. pseudonana genome encodes six proteins
which are most closely related to genes encoded by the Guillardia
theta (cryptomonad) nucleomorph genome. Four of these genes are
also found in red algal plastid genomes, thus demonstrating
successive EGT from red algal plastid to red algal nucleus
(nucleomorph) to heterokont host nucleus.[139] More recent
phylogenomic analyses of diatom proteomes provided evidence for
a prasinophyte-like endosymbiont in the common ancestor of
chromalveolates as supported by the fact the 70% of diatom genes
of Plantae origin are of green lineage provenance and that such
genes are also found in the genome of other stramenopiles.
Therefore, it was proposed that chromalveolates are the product of
serial secondary endosymbiosis first with a green algae, followed by
a second one with a red algae that conserved the genomic
footprints of the previous but displaced the green plastid.[143]
However, phylogenomic analyses of diatom proteomes and
chromalveolate evolutionary history will likely take advantage of
complementary genomic data from under-sequenced lineages such
as red algae.
Horizontal gene transfer
In addition to EGT, horizontal gene transfer (HGT) can occur

independently of an endosymbiotic event. The publication of the P.
tricornutum genome reported that at least 587 P. tricornutum genes
appear to be most closely related to bacterial genes, accounting for
more than 5% of the P. tricornutum proteome. About half of these are
also found in the T. pseudonana genome, attesting their ancient
incorporation in the diatom lineage.[140]
Genetic engineering
To understand the biological mechanisms which underlie the great

importance of diatoms in geochemical cycles, scientists have used
the Phaeodactylum tricornutum and Thalassiosira spp. species as
model organisms since the 90's.[144] Few molecular biology tools are
currently available to generate mutants or transgenic lines :
plasmids containing transgenes are inserted into the cells using the
biolistic method[145] or transkingdom bacterial conjugation[146] (with
10−6 and 10−4 yield respectively[145][146]), and other classical
transfection methods such as electroporation or use of PEG have
been reported to provide results with lower efficiencies.[146]
Transfected plasmids can be either randomly integrated into the

diatom's chromosomes or maintained as stable circular episomes
(thanks to the CEN6-ARSH4-HIS3 yeast centromeric sequence[146]).
The phleomycin/zeocin resistance gene Sh Ble is commonly used
as a selection marker,[144][147] and various transgenes have been
successfully introduced and expressed in diatoms with stable
transmissions through generations,[146][147] or with the possibility to
remove it.[147]
Furthermore, these systems now allow the use of the CRISPR-Cas
genome edition tool, leading to a fast production of functional
knock-out mutants[147][148] and a more accurate comprehension of
the diatoms' cellular processes.
Human uses
P
a
l
e
o
n
t Diatomaceous earth consisting of centric (radially symmetric) and
o pennate (bilaterally symmetric) diatoms suspended in water.
l (click 3 times to fully enlarge)
o
g
y
Decomposition and decay of diatoms leads to organic and inorganic

(in the form of silicates) sediment, the inorganic component of
which can lead to a method of analyzing past marine environments
by corings of ocean floors or bay muds, since the inorganic matter is
embedded in deposition of clays and silts and forms a permanent
geological record of such marine strata (see siliceous ooze).
Industrial
Diatoms, and their shells (frustules) as diatomite or diatomaceous

earth, are important industrial resources used for fine polishing and
liquid filtration. The complex structure of their microscopic shells
has been proposed as a material for nanotechnology.[149]
Diatomite is considered to be a natural nano material and has many

uses and applications such as: production of various ceramic
products, construction ceramics, refractory ceramics, special oxide
ceramics, for production of humidity control materials, used as
filtration material, material in the cement production industry, initial
material for production of prolonged-release drug carriers,
absorption material in an industrial scale, production of porous
ceramics, glass industry, used as catalyst support, as a filler in
plastics and paints, purification of industrial waters, pesticide holder,
as well as for improving the physical and chemical characteristics of
certain soils, and other uses.[150][151][152]
Diatoms are also used to help determine the origin of materials

containing them, including seawater.
Nanotechnology
The deposition of silica by diatoms may also prove to be of utility to

nanotechnology.[153] Diatom cells repeatedly and reliably
manufacture valves of various shapes and sizes, potentially allowing
diatoms to manufacture micro- or nano-scale structures which may
be of use in a range of devices, including: optical systems;
semiconductor nanolithography; and even vehicles for drug delivery.
With an appropriate artificial selection procedure, diatoms that
produce valves of particular shapes and sizes might be evolved for
cultivation in chemostat cultures to mass-produce nanoscale
components.[154] It has also been proposed that diatoms could be
used as a component of solar cells by substituting photosensitive
titanium dioxide for the silicon dioxide that diatoms normally use to
create their cell walls.[155] Diatom biofuel producing solar panels
have also been proposed.[156]
Supporting and regulating services provided by marine diatoms and so

of their negative impacts
CNN = cloud
condensation nuclei, DMS = dimethylsulphide, DMSP =
dimethylsulfoniopropionate, VOCs = volatile organic
compounds
dashe
arrow: negative effect, solid arrow: positive effects
Forensic
The main goal of diatom analysis in forensics is to differentiate a

death by submersion from a post-mortem immersion of a body in
water. Laboratory tests may reveal the presence of diatoms in the
body. Since the silica-based skeletons of diatoms do not readily
decay, they can sometimes be detected even in heavily decomposed
bodies. As they do not occur naturally in the body, if laboratory tests
show diatoms in the corpse that are of the same species found in
the water where the body was recovered, then it may be good
evidence of drowning as the cause of death. The blend of diatom
species found in a corpse may be the same or different from the
surrounding water, indicating whether the victim drowned in the
same site in which the body was found.[157]
History of discovery
Tabellaria is a genus of freshwater

diatoms, cuboid in shape with
frustules (siliceous cell walls)
attached at the corners so the
colonies assume a zigzag shape.
The first illustrations of diatoms are found in an article from 1703 in

Transactions of the Royal Society showing unmistakable drawings
of Tabellaria (https://academic.oup.com/view-large/figure/21455463
3/fbaa049f4.tif) .[158] Although the publication was authored by an
unnamed English gentleman, there is recent evidence that he was
Charles King of Staffordshire.[158][159] It is only 80 years later that we
find the first formally identified diatom, the colonial Bacillaria
paxillifera, discovered and described in 1783 by Danish naturalist
Otto Friedrich Müller.[158] Like many others after him, he wrongly
thought that it was an animal due to its ability to move. Even Charles
Darwin saw diatom remains in dust whilst in the Cape Verde Islands,
although he was not sure what they were. It was only later that they
were identified for him as siliceous polygastrics. The infusoria that
Darwin later noted in the face paint of Fueguinos, native inhabitants
of Tierra del Fuego in the southern end of South America, were later
identified in the same way. During his lifetime, the siliceous
polygastrics were clarified as belonging to the Diatomaceae, and
Darwin struggled to understand the reasons underpinning their
beauty. He exchanged opinions with the noted cryptogamist G. H. K.
Thwaites on the topic. In the fourth edition of On the Origin of
Species he stated that "Few objects are more beautiful than the
minute siliceous cases of the diatomaceae: were these created that
they might be examined and admired under the high powers of the
microscope"? and reasoned that their exquisite morphologies must
have functional underpinnings rather than having been created
purely for humans to admire.[160]
See also
Highly branched isoprenoid, long-chain alkenes produced by a
small number of marine diatoms
Notes
a. From Greek: διατομή, romanized: diatomé, "a cutting through, a
severance",[6] from Greek: διάτομος, romanized: diátomos, "cut in half,
divided equally" [7] from Greek: διατέμνω, romanized: diatémno, "to cut in
twain".[8][9]: 718
References
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External links
Wikispecies has information related to Diatoms.
Wikimedia Commons has media related to Diatoms.
Diatom EST database (http://www.biologie.ens.fr/diatomics/ES
T/) , École Normale Supérieure
Plankton*Net (http://planktonnet.awi.de/) , taxonomic database
including images of diatom species
Life History and Ecology of Diatoms (http://www.ucmp.berkeley.e
du/chromista/diatoms/diatomlh.html) , University of California
Museum of Paleontology
Diatoms: 'Nature's Marbles' (https://web.archive.org/web/200702
09042756/http://hjs.geol.uib.no/Diatoms/index.html-ssi) , Eureka
site, University of Bergen
Diatom life history and ecology (http://www.ucl.ac.uk/GeolSci/mic
ropal/diatom.html) , Microfossil Image Recovery and Circulation
for Learning and Education (MIRACLE), University College London
Diatom page (http://www.rbge.org.uk/science/cryptogamic-plants
-and-fungi/phycology/diatoms) Archived (https://web.archive.or
g/web/20091008014752/http://www.rbge.org.uk/science/cryptog
amic-plants-and-fungi/phycology/diatoms) 8 October 2009 at the
Wayback Machine, Royal Botanic Garden Edinburgh
Geometry and Pattern in Nature 3: The holes in radiolarian and
diatom tests (http://www.microscopy-uk.org.uk/mag/artfeb05/cb
diatoms.html)
Diatom QuickFacts (https://web.archive.org/web/2008052320303
5/http://www.mbari.org/staff/conn/botany/phytoplankton/phytop
lankton_diatoms.htm) , Monterey Bay Aquarium Research
Institute
Algae image database (http://diatom.ansp.org/algae_image/)
Academy of Natural Sciences of Philadelphia (ANSP)
Diatom taxa (http://diatom.ansp.org/nawqa/Taxalist.aspx)
Academy of Natural Sciences of Philadelphia (ANSP)
An Introduction to the Microscopical Study of Diatoms (http://ww
w.microscopy-uk.org.uk/diatomist/rbm_US_Royal.pdf) by Robert
B. McLaughlin
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Diatom

A diatom (Neo-Latin diatoma)[a] is any member of a large group

Light microscopy of a sampling of marine diatoms found living between

Diatomea Dumortier, 1821[2]

Diatoms are unicellular organisms: they occur either as solitary cells

Dinotoms are diatoms that have become endosymbionts inside

The study of diatoms is a branch of phycology. Diatoms are

Fossil evidence suggests that diatoms originated during or before

Diatoms are used to monitor past and present environmental

Dwindling diatoms and the mixed layer

3D-animation of the diatom Corethron sp.

The cell itself consists of two halves, each containing an essentially

2. Nucleolus; location of the chromosomes

3. Golgi apparatus; modifies proteins and sends them out of

4. Cell wall; outer membrane of the cell

5. Pyrenoid; center of carbon fixation

6. Chromatophore; pigment carrying membrane structure

7. Vacuoles; vesicle of a cell that contains fluid bound by a

8. Cytoplasmic strands; hold the nucleus

9. Mitochondria; create ATP (energy) for the cell

b. Striae (pores, punctae, spots or dots in a line on the surface)

c. Raphe (slit in the valves)

d. Central nodule (thickening of wall at the midpoint of raphe)

e. Stigmata (holes through valve surface which looks rounded

f. Punctae (spots or small perforations on the surface)

g. Polar nodules (thickening of wall at the distal ends of the

Selections from Ernst Haeckel's 1904

Pennate diatoms are bilaterally symmetric. Each one of their valves

Shape classification of diatom frustules. The

Structure of a centric diatom frustule [37]

Pennate diatom from an Arctic meltpond, infected

The exact mechanism of transferring silica absorbed by the diatom

These silica transport proteins are unique to diatoms, with no

The first characterisations of the biochemical processes and

Chaetoceros willei Chaetoceros furcillatus J.W.Bailey,

Some pennate diatoms are capable of a type of locomotion called

Planktonic diatoms such as Thalassiosira sp. (56-62),

Some Thalassiosira diatoms form chain-like colonies, like these

Centric diatom (oogamy) Pennate diatom (morphological

Reproduction and cell size

Reproduction among these organisms is asexual by binary fission,

A defining characteristic of all diatoms is their restrictive and

Diatoms are mostly non-motile; however, sperm found in some

Certain species of bacteria in oceans and lakes can accelerate the

Regions of high abundance of diatoms in the ocean

Diatom dominance (as a percentage of total cell counts)

Diatoms are a widespread group and can be found in the oceans, in

Planktonic diatoms in freshwater and marine environments typically

The freshwater diatom Didymosphenia geminata, commonly known

Ultimately, diatom cells in these resting populations re-enter the

In the open ocean, the diatom (spring) bloom is typically ended by a

Because of this bloom-and-bust cycle, diatoms are believed to play a

Reason for success

Diatoms are ecologically successful, and occur in virtually every

Sources for collection

Diatoms can be obtained from multiple sources.[78] Marine diatoms

However, the precise timing of the "take-over" remains unclear, and

The diagram depicts some mechanisms by which marine diatoms

While often overlooked in photosynthetic organisms, the

Diatoms are mainly photosynthetic; however a few are obligate

Photosynthetic diatoms that find themselves in an environment

Major pigments of diatoms are chlorophylls a and c, beta-carotene,

Light microscopy of several species