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Ocean's Crystalline Architects

Delve into the fascinating world of Acantharia, unique marine microplankton with intricate strontium sulfate skeletons, their biology, ecology, and symbiotic relationships.

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Acantharia: An Overview

Microscopic Marine Wonders

Acantharia represent a distinct class of radiolarian protozoa, single-celled organisms that are a vital component of marine microplankton. Their most striking feature is their intricate internal skeleton, uniquely composed of strontium sulfate. These organisms typically range in size from approximately 200 micrometers to several millimeters, making them visible under a microscope and, in some cases, to the naked eye.

Heterotrophs and Mixotrophs

Primarily, Acantharia are heterotrophic, meaning they obtain nutrients by consuming other organisms. However, a significant number of acantharian species engage in a fascinating symbiotic relationship with photosynthetic endosymbionts. This allows them to function as mixotrophs, deriving energy from both predation and photosynthesis, a strategy that is particularly advantageous in nutrient-poor oceanic environments.

Ecological Significance

As marine microplankton, Acantharia play a role in the oceanic food web and biogeochemical cycles. Their unique strontium sulfate skeletons, while not fossilizing, contribute to the vertical flux of organic carbon in the ocean. The rapid sedimentation of their cysts, particularly observed in regions like the Iceland Basin and the Southern Ocean, highlights their contribution to the deep ocean's particle flux.

Intricate Morphology

The Strontium Sulfate Skeleton

The defining characteristic of Acantharia is their skeleton, meticulously constructed from strontium sulfate (SrSO4) in the form of celestine crystals. This biomineralization of strontium sulfate is a unique capability among marine organisms. Celestine, known for its delicate blue hue and significant density, acts as a mineral ballast, facilitating the rapid sinking of acantharian remains and cysts to bathypelagic depths upon the organism's demise. Unlike the silica skeletons of other radiolarians, acantharian skeletons do not fossilize due to the scarcity of strontium sulfate in seawater and the subsequent dissolution of the crystals post-mortem.

Precise Spine Architecture

The arrangement of the acantharian spines is remarkably precise, adhering to a geometric principle known as the Müllerian law. This law describes the spines as lying at the intersections of five lines of latitude, symmetrically positioned around an equator, and eight lines of longitude, uniformly spaced. Each longitudinal line alternately bears either two "tropical" spines or one "equatorial" and two "polar" spines, creating a highly ordered and aesthetically complex skeletal framework.

Dual Cytoplasmic Regions

The cellular structure of Acantharia is distinctly divided into two primary regions: the endoplasm and the ectoplasm. The endoplasm, forming the central core of the cell, houses the main organelles and numerous nuclei. It is separated from the ectoplasm by a capsular wall, a mesh-like structure composed of microfibrils. In species that host symbiotic algae, these algal partners are typically found within the endoplasm. The ectoplasm, extending outwards, is crucial for prey capture and contains food vacuoles for digestion. Surrounding the ectoplasm is a periplasmic cortex, also made of microfibrils, arranged into twenty distinct plates, each perforated to allow a spicule to project through. Contractile myonemes connect this cortex to the spines, enabling the ectoplasm to expand and contract, thereby regulating the cell's buoyancy.

Classification & Phylogeny

Spine Junctions as Classifiers

The primary method for classifying Acantharia hinges on the manner in which their skeletal spines converge at the cell's center. Skeletons are composed of either ten diametric spicules, which traverse the cell's core, or twenty radial spicules, which terminate at the center. The nature of this central junction—whether tight or flexible—is a key distinguishing feature. Acantharians possessing diametric spicules or loosely attached radial spicules exhibit the remarkable ability to rearrange or even shed their spicules, a mechanism often associated with the formation of protective cysts.

Orders of Acantharia

The class Acantharia is traditionally divided into several orders based on these skeletal characteristics. While morphological classifications generally align with phylogenetic trees derived from ribosomal RNA gene sequences, it is noted that many of these groups are polyphyletic, indicating complex evolutionary pathways.

The primary orders of Acantharia include:

  • Holacanthida: Characterized by 10 diametric spicules that simply cross without a central junction. These species are capable of encystment and are considered to have evolved earlier, encompassing molecular clades A, B, and D.
  • Chaunacanthida: Possess 20 radial spicules that are loosely attached at the cell's center. Like Holacanthida, they can form cysts and represent an intermediate evolutionary stage, corresponding to molecular clade C.
  • Symphyacanthida: Distinguished by 20 radial spicules that form a tight central junction. These are among the more recently evolved groups, constituting parts of molecular clades E and F, and typically feature more complex skeletons.
  • Arthracanthida: Also exhibit 20 radial spines with a tight central junction, sharing the more recent evolutionary trajectory and complex skeletal structures with Symphyacanthida, belonging to molecular clades E and F.

This classification system provides a framework for understanding the diversity within Acantharia, reflecting both their structural variations and evolutionary relationships.

The Symbiotic Life

Photosymbiosis in the Endoplasm

A remarkable feature of many acantharians is their engagement in photosymbiosis, hosting single-celled algae within their inner cytoplasm, the endoplasm. This includes certain species within clade B (Holacanthida) and all species in clades E and F (Symphiacanthida and Arthracanthida). This symbiotic relationship transforms these acantharians into mixotrophs, enabling them to harness energy through both heterotrophic feeding and autotrophic photosynthesis. This dual energy acquisition strategy is hypothesized to be crucial for their prevalence in the oligotrophic (low-nutrient) regions of the oceans and may also provide the substantial energy required to maintain their elaborate strontium sulfate skeletons.

A Reciprocal Exchange?

The symbiotic interaction is thought to involve a reciprocal exchange of resources. It is hypothesized that the acantharians supply their algal partners with essential nutrients, such as nitrogen and phosphorus, which they obtain through the capture and digestion of prey. In return, the algae provide the acantharians with sugars produced during photosynthesis. However, the precise nature of the benefit to the algal symbionts remains an area of active research; it is not definitively known whether the algae truly benefit from the relationship or if they are primarily exploited and subsequently digested by their acantharian hosts.

Diverse Symbiont Communities

The diversity of algal symbionts varies across different acantharian clades. Symbiotic Holacanthida acantharians are known to host a wide array of symbiont assemblages, including several genera of dinoflagellates (e.g., Pelagodinium, Heterocapsa, Scrippsiella, Azadinium) and a haptophyte (Chrysochromulina). In contrast, acantharians from clades E and F exhibit a more specific symbiosis, predominantly hosting symbionts from the haptophyte genus Phaeocystis, though Chrysochromulina symbionts are occasionally observed. Intriguingly, clade F acantharians can simultaneously host multiple species and strains of Phaeocystis. The observed discrepancy between the internal symbiont community and the external availability of potential symbionts suggests that acantharians are selective in their choice of partners and likely maintain these symbionts for extended periods rather than continuously digesting and recruiting new ones.

The Enigmatic Lifecycle

Reproduction and Encystment

Acantharian adults are typically multinucleated organisms. A key aspect of their reproductive strategy, particularly in earlier diverging clades, involves the ability to shed their characteristic spines and form cysts. These structures are often referred to as reproductive cysts, indicating their role in the organism's life cycle. Reproduction is believed to occur through the formation of "swarmer cells," which are often flagellate. Observations in laboratory settings have shown both cysts and non-encysted cells releasing these swarmers. However, a complete understanding of the acantharian life cycle remains elusive, as the fusion of swarmers to produce a new acantharian has yet to be directly observed.

Deep-Water Dispersal Strategy

Acantharian cysts are frequently detected in sediment traps, leading to the hypothesis that these cysts play a crucial role in enabling acantharians to sink into the deep ocean. This deep-water dispersal strategy is thought to enhance the survival prospects of juvenile acantharians by releasing swarmer cells in environments potentially more conducive to their development. Furthermore, genetic data combined with imaging studies suggest that even non-cyst-forming acantharians may undertake a similar descent to deeper waters for the release of their swarmers. The inability to sustain these organisms through successive generations in laboratory cultures has significantly hindered comprehensive research into their full life cycle.

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References

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