# Desiccation Tolerance as the Basis of Long-Term Seed Viability

**Authors:** Galina Smolikova, Tatiana Leonova, Natalia Vashurina, Andrej Frolov, Sergei Medvedev

PMC · DOI: 10.3390/ijms22010101 · International Journal of Molecular Sciences · 2020-12-24

## TL;DR

This paper explains how seeds can survive long periods of dryness due to specific genes and molecular mechanisms activated during maturation.

## Contribution

The paper highlights the evolution of desiccation tolerance mechanisms in plants and their regulation during seed development.

## Key findings

- Orthodox seeds can lose up to 95% of their water and remain viable due to desiccation tolerance.
- LEA proteins, sHSP, oligosaccharides, and antioxidants are key to desiccation tolerance in seeds.
- Abscisic acid and DOG1 regulate a network of transcription factors and epigenetic mechanisms for seed maturation.

## Abstract

Desiccation tolerance appeared as the key adaptation feature of photoautotrophic organisms for survival in terrestrial habitats. During the further evolution, vascular plants developed complex anatomy structures and molecular mechanisms to maintain the hydrated state of cell environment and sustain dehydration. However, the role of the genes encoding the mechanisms behind this adaptive feature of terrestrial plants changed with their evolution. Thus, in higher vascular plants it is restricted to protection of spores, seeds and pollen from dehydration, whereas the mature vegetative stages became sensitive to desiccation. During maturation, orthodox seeds lose up to 95% of water and successfully enter dormancy. This feature allows seeds maintaining their viability even under strongly fluctuating environmental conditions. The mechanisms behind the desiccation tolerance are activated at the late seed maturation stage and are associated with the accumulation of late embryogenesis abundant (LEA) proteins, small heat shock proteins (sHSP), non-reducing oligosaccharides, and antioxidants of different chemical nature. The main regulators of maturation and desiccation tolerance are abscisic acid and protein DOG1, which control the network of transcription factors, represented by LEC1, LEC2, FUS3, ABI3, ABI5, AGL67, PLATZ1, PLATZ2. This network is complemented by epigenetic regulation of gene expression via methylation of DNA, post-translational modifications of histones and chromatin remodeling. These fine regulatory mechanisms allow orthodox seeds maintaining desiccation tolerance during the whole period of germination up to the stage of radicle protrusion. This time point, in which seeds lose desiccation tolerance, is critical for the whole process of seed development.

## Linked entities

- **Genes:** LEA (dehydrin LEA) [NCBI Gene 816688], LOC8061347 (16.9 kDa class I heat shock protein 3) [NCBI Gene 8061347], ADGRL1 (adhesion G protein-coupled receptor L1) [NCBI Gene 22859], FUS3 (AP2/B3-like transcriptional factor family protein) [NCBI Gene 822293], ABI3 (ABI family member 3) [NCBI Gene 51225], ABI5 (Basic-leucine zipper (bZIP) transcription factor family protein) [NCBI Gene 818199], AGL67 (AGAMOUS-like 67) [NCBI Gene 844130], LOC100247272 (protein RGF1 INDUCIBLE TRANSCRIPTION FACTOR 1) [NCBI Gene 100247272], ANO1 (anoctamin 1) [NCBI Gene 55107]
- **Proteins:** LOC8061347 (16.9 kDa class I heat shock protein 3), ANO1 (anoctamin 1)
- **Chemicals:** abscisic acid (PubChem CID 30583)

## Full-text entities

- **Genes:** PROS1 (protein S) [NCBI Gene 478529], ANO1 (anoctamin 1) [NCBI Gene 55107] {aka DOG1, INDMS, MYMY7, ORAOV2, TAOS2, TMEM16A}, ABI3 (ABI family member 3) [NCBI Gene 51225] {aka NESH, SSH3BP3}, beta-1,3-Glucanase [NCBI Gene 107784423], PIMT1 (protein-l-isoaspartate methyltransferase 1) [NCBI Gene 823991] {aka ATPIMT1, Arabidopsis thaliana protein-l-isoaspartate methyltransferase 1, protein-l-isoaspartate methyltransferase 1}, LEC1 (Histone superfamily protein) [NCBI Gene 838800] {aka AtLEC1, EMB 212, EMB212, EMBRYO DEFECTIVE 212, LEAFY COTYLEDON 1, NF-YB9}, SAG21 (senescence-associated gene 21) [NCBI Gene 828053] {aka Arabidopsis thaliana late embryogenensis abundant like 5, AtLEA5, T14P8.2, T14P8_2, senescence-associated gene 21}, HSP70 (heat shock protein 70) [NCBI Gene 403612] {aka HSPA1}, AGL67 (AGAMOUS-like 67) [NCBI Gene 844130] {aka AGAMOUS-like 67, F28K19.16, F28K19_16}, LPP2 (lipid phosphate phosphatase 2) [NCBI Gene 838072] {aka ATLPP2, ATPAP2, F9L1.2, F9L1_2, LIPID PHOSPHATE PHOSPHATASE 2, PHOSPHATIDIC ACID PHOSPHATASE 2}, ADGRL1 (adhesion G protein-coupled receptor L1) [NCBI Gene 22859] {aka CIRL1, CL1, DEDBANP, LEC2, LPHN1}, LEC2 (AP2/B3-like transcriptional factor family protein) [NCBI Gene 839724] {aka AtLEC2, F3H9.5, F3H9_5, LEAFY COTYLEDON 2}, AT3G24640 (lyase) [NCBI Gene 822060], DOG1 (delay of germination 1) [NCBI Gene 834623] {aka ATDOG1, DELAY OF GERMINATION 1, GAAS5, GLUCOSE SENSING QTL 5, GSQ5, K15I22.3}, ABI2 (abl interactor 2) [NCBI Gene 488485], PNLIPRP1 (pancreatic lipase related protein 1) [NCBI Gene 404010] {aka PLRP1}, ABI1 (abl interactor 1) [NCBI Gene 607247], LEA14 (Late embryogenesis abundant protein) [NCBI Gene 837071] {aka F22L4.3, F22L4_3, LATE EMBRYOGENESIS ABUNDANT 14, LEA1, LIGHT STRESS-REGULATED 3, LSR3}, ABI5 (Basic-leucine zipper (bZIP) transcription factor family protein) [NCBI Gene 818199] {aka ABA INSENSITIVE 5, AtABI5, F2H17.12, F2H17_12, GIA1, GROWTH-INSENSITIVITY TO ABA 1}, FUS3 (AP2/B3-like transcriptional factor family protein) [NCBI Gene 822293] {aka FUSCA 3}, LEA3 (Late embryogenesis abundant 3 (LEA3) family protein) [NCBI Gene 839304] {aka AtLEA3, F22D16.18, F22D16_18, late embryogenesis abundant 3}, CAT (catalase) [NCBI Gene 403474], ABA1 (zeaxanthin epoxidase (ZEP) (ABA1)) [NCBI Gene 836838] {aka ABA DEFICIENT 1, ARABIDOPSIS THALIANA ABA DEFICIENT 1, ARABIDOPSIS THALIANA ZEAXANTHIN EPOXIDASE, ATABA1, ATZEP, IBS3}, PARP1 (poly(ADP-ribose) polymerase 1) [NCBI Gene 142] {aka ADPRT, ADPRT 1, ADPRT1, ARTD1, PARP, PARP-1}, MIR172a (ncRNA) [NCBI Gene 28718310] {aka MICRORNA 172, MIR172, microRNA172A, p_MI0000215}, HSFA9 (heat shock transcription factor A9) [NCBI Gene 835493] {aka ARABIDOPSIS THALIANA HEAT SHOCK TRANSCRIPTION FACTOR  A9, ARABIDOPSIS THALIANA HEAT SHOCK TRANSCRIPTION FACTOR A9, AT-HSFA9, MJP23.4, MJP23_4, heat shock transcription factor  A9}, HSPD1 (heat shock protein family D (Hsp60) member 1) [NCBI Gene 478854] {aka CPN60, HSP-60, Hsp60}, ALB (albumin) [NCBI Gene 403550] {aka CSA}, ABI3 (AP2/B3-like transcriptional factor family protein) [NCBI Gene 822061] {aka ABA INSENSITIVE 3, ABSCISIC ACID INSENSITIVE 3, AtABI3, SIS10, SUGAR INSENSITIVE 10}, PRC1 (protein regulator of cytokinesis 1) [NCBI Gene 488742], ADGRL2 (adhesion G protein-coupled receptor L2) [NCBI Gene 23266] {aka CIRL2, CL2, LEC1, LPHH1, LPHN2}
- **Diseases:** AGEs (MESH:D003643), dehydration (MESH:D003681), AGE (OMIM:613784), AR (MESH:D000094025), seed damage (MESH:D009366), water loss (MESH:D000069578)
- **Chemicals:** nitrates (MESH:D009566), disulfide (MESH:D004220), peroxynitrite (MESH:D030421), beta-glucans (MESH:D047071), NAD+ (MESH:D009243), Glu (MESH:D018698), alcohols (MESH:D000438), aspartate (MESH:D001224), dimethyl sulfoxide (MESH:D004121), salt (MESH:D012492), glucose (MESH:D005947), Chlorophylls (MESH:D002734), RNS (MESH:D026361), thiourea (MESH:D013890), glutathione disulfide (MESH:D019803), Tocopherols (MESH:D024505), fatty acid (MESH:D005227), oxygen (MESH:D010100), 5'-methylcytosine (MESH:D044503), poly (ADP-ribose) (MESH:D011064), phospholipids (MESH:D010743), glyoxal (MESH:D006037), carotenoids (MESH:D002338), raffinose (MESH:D011887), superoxide (MESH:D013481), Base (MESH:D009711), methionine (MESH:D008715), H2O2 (MESH:D006861), deoxyribose (MESH:D003855), fats (MESH:D005223), mannitol (MESH:D008353), water (MESH:D014867), hydroxyl radical (MESH:D017665), 1O2 (-), 3-deoxyglucosone (MESH:C016350), GA (MESH:D005708), Gibberellic acid (MESH:C007842), fructose (MESH:D005632), sorbitol (MESH:D013012), Carbohydrates (MESH:D002241), nitrogen (MESH:D009584), ABA (MESH:D000040), L-amino acid (MESH:D000596), sucrose (MESH:D013395), oligosaccharides (MESH:D009844), Thr (MESH:D013912), singlet oxygen (MESH:D026082), heme (MESH:D006418), oils (MESH:D009821), Gly (MESH:D005998), ADP-ribose (MESH:D000246), isoaspartic acid (MESH:D026581), Glutathione (MESH:D005978), methionine sulfoxide (MESH:C013111), ethylene (MESH:C036216), methylglyoxal (MESH:D011765), tocotrienols (MESH:D024508), GA1 (MESH:C011258), asparagine (MESH:D001216), guanine (MESH:D006147)
- **Species:** Powellomyces sp. EA (species) [taxon 252690], Medicago sativa (alfalfa, species) [taxon 3879], PX clade (clade) [taxon 569578], Solanum tuberosum (potatoes, species) [taxon 4113], Nicotiana tabacum (American tobacco, species) [taxon 4097], Homo sapiens (human, species) [taxon 9606], Acer pseudoplatanus (sycamore maple, species) [taxon 4026], Phoenix dactylifera (date palm, species) [taxon 42345], Pyrus communis (pear, species) [taxon 23211], Glycine max (soybean, species) [taxon 3847], Helianthus annuus (common sunflower, species) [taxon 4232], Lactuca sativa (cultivated lettuce, species) [taxon 4236], Ricinus communis (castor bean, species) [taxon 3988], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Camellia sinensis (black tea, species) [taxon 4442], Oryza sativa (Asian cultivated rice, species) [taxon 4530], Lathyrus oleraceus (garden pea, species) [taxon 3888]
- **Cell lines:** abi3-5 — Homo sapiens (Human), Induced pluripotent stem cell (CVCL_C6TI)

## Full text

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## Figures

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## References

177 references — full list in the complete paper: https://tomesphere.com/paper/PMC7795748/full.md

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Source: https://tomesphere.com/paper/PMC7795748