Unit – II - ANATOMY

TOPIC: P- PROTEINS

By –
Sonakshi Srivastava
Post- Graduate Botany Student
Department of Botany
Gujarat University, Ahmedabad
 P - Proteins
• What are P – proteins?
• P – proteins are a category of structural, main phloem proteins, present in angiosperms (all eudicots and many
monocots), involved in sealing damaged elements so as to avoid leaking of sieve element sap, which is rich in sugars
and other organic molecules.
• However, they are absent in pteridophytes and gymnosperms.
• In earlier scientific literature, they were referred to as slime.
• P- protein occurs in several different forms (Tubular, Fibrillar, Granular, and Crystalline), depending upon the species
of angiosperm and the maturity of the cell.
Eg: In immature cells, P- proteins are evident as discrete bodies in the cytosol, known as P- protein bodies, which may be
Spheroidal, Spindle-shaped, or Twisted and Coiled.
They often disperse to Tubular or Fibrillar forms during cell maturation.
• In a completely mature sieve tube element, the P- proteins are located in the periphery region of the cytosol.
• Dictyosomes are reported to be involved in the synthesis of P- proteins.
Chapter - 11:
Translocation in Phloem

Pg - 287

Plant Physiology and

Development (6th Edition)
by Taiz and Zeiger
 Function and action of mechanism of P- Proteins
(Part – I)
• P-protein appears to function in sealing off damaged sieve elements by plugging sieve plate pores.
• Sieve tubes are under very high internal turgor pressure, and the sieve elements in a sieve tube are connected through
sieve plate pores that appear to be open.
• When a sieve tube is cut or punctured, the release of pressure causes the contents of the sieve elements to surge toward
the cut end, from which the plant could lose much sugar-rich phloem sap if there were no sealing mechanism.
• When surging occurs, however, P-protein is trapped on the sieve plate pores, helping to seal the sieve element and
prevent further loss of sap.
• Direct support for the sealing function of P-protein has been found in both tobacco and Arabidopsis, in which mutants
lacking P-protein lose significantly more transport sugar by sap exudation after wounding than do wild-type plants.
• However, no visible phenotypic differences were observed between the mutant and wild-type plants.
 (Part – II)
• Protein crystals released from ruptured plastids may play a similar sealing role in some monocots as P-protein does in
eudicots.
• The sieve element organelles (mitochondria, plastids, and ER), however, appear to be anchored to each other or to the
sieve element plasma membrane by minute protein “clamps.” Which organelles are anchored depends on the species.
• Another mechanism for blocking wounded sieve tubes with proteins occurs in plants in the legume family (Fabaceae).
These plants contain large crystalloid P-protein that do not disperse during development.
• However, following damage or osmotic shock, the P-protein rapidly disperse and block the sieve tube. The process is
reversible and controlled by calcium ions (Ca2+). These P-protein, known as Forisomes, occur only in certain legumes
and are encoded by members of the Sieve Element Occlusion (SEO) gene family.
• Homologous members of the SEO gene family encode conventional P-proteins in other species. These are called SEOR
(Sieve Element Occlusion Related) genes.
 (Part – III)
• The term P-protein thus includes similar molecules that are involved in blocking wounded sieve elements in all eudicot
angiosperms, as well as special P-proteins, such as forisomes and PP1 (P-Protein 1) and PP2 (P-Protein 2), found in
Cucurbita maxima (Winter squash).
• Both PP1 and PP2 are synthesized in the companion cells and transported to the sieve tube element.
• PP1 is present in all the developmental stages of the sieve tubes, however, PP2 is produced only after the complete
maturation of the sieve tubes.
 (Part – IV)
• An Additional Mechanism for Blocking Wounded Sieve Elements in the Legume Family
Susan Dunford, University of Cincinnati
• While other plant families have protein bodies that do not disperse during development, it is unknown whether other
plants have similar mechanisms for plugging wounded sieve tubes.
• Since callose synthesis is also calcium-dependent, wounding in Vicia faba (broad bean) further causes callose
deposition in sieve pores following protein-body dispersal (Furch et al. 2007).
 Classification of P- proteins

• Non-Dispersive P-protein bodies

 The Non-Dispersive P-protein bodies are formed early during ontogeny (ontogeny = the development of
individual cells or organisms) in young sieve tubes.
 Once the sieve element matures, the sieve pores open, and they become active in translocation; non-dispersive
P-protein bodies remain as visible large bodies within the sieve tube.
 One of the examples of these proteins is Forisomes which occur in faboid legumes (bean plants) only.
 They undergo a rapid, reversible switch from an elongated low-volume conformation into a spherical high-
volume conformation.
 The high-volume state is induced when Ca2+ enters the cytoplasm of disturbed or severed sieve elements.
High volume-state forisomes impede flow in the sieve tube.
 About 10% of the Angiosperm species contain non-dispersive proteins.
Non-dispersive P-protein bodies (asterisk) as seen in the confocal
microscope (left) and the transmission electron microscope
(right). SE= sieve element, CC = companion cell.

From Knoblauch et al. (2001) Calcium regulated stopcocks in

legume sieve tubes. Plant Cell 13, 1221 – 1230.

Copyright American Society of Plant Biologists

 Classification of P- proteins

• Dispersive P-protein bodies

 The Dispersive P-protein bodies are also formed early during ontogeny in young sieve tubes.
 However, once the sieve element matures, the bodies do not remain in the lumen of the tube, but disperse into
fine filaments.
 The rest, 90% of the Angiosperm species possess non-dispersive elements.
A bundle of filaments (left) and individual filaments (right) in mature
sieve elements, derived from dispersive P-protein bodies.

From Froelich et al. (2011) Phloem ultrastructure and pressure flow: SEOR
protein agglomerations do not affect translocation. Plant Cell
doi/10.1105/tpc.111.093179

Copyright American Society of Plant Biologists

 References

 Plant Physiology and Development (6th Edition) - Eduardo Zeiger, Lincoln Taiz, Ian Max Moller, Angus
Murphy
 Michael Knoblauch Lab – Washington State University
 P- proteins: Structure, Classification and Functions – Easy Biology Class
 For more references, kindly refer to the research papers.