Suaeda salsa

Suaeda salsa (L.) Pall.
Flora ID 2318·Amaranthaceae (older literature may use Chenopodiaceae)·Suaeda

Flora account

Overview

Suaeda salsa occurs in both intertidal and inland saline habitats in the Yellow River Delta. Controlled studies cover seeds, seedlings, established plants, and reproduction; a 200 mM NaCl pot study continued to mature seed, while higher-concentration results are interpreted by life stage and endpoint. [S13, S01, S02, S03, S11]

Name and taxonomy

Floras and the relevant primary studies use Suaeda salsa (L.) Pall.; seed and seedling results identify intertidal and inland sources separately. [S13]

Morphology and identification

Flora of China describes an annual herb 20–80 cm tall, with an erect stem branching mainly above. Linear semiterete leaves are usually 1–2.5 cm long; plants may be green or purple-red. Axillary flower clusters form interrupted spikes along branches. [S13]

Distribution, habitat, and phenology

The Yellow River Delta has both intertidal and inland saline populations. A 2021 field study reported total soil salts of 11.92±1.59‰ at intertidal sites and 3.84±1.63‰ above the tide line. A 2020 study also sampled plants and root-associated soil from both habitat types. [S01, S02, S03, S11]

Salinity in natural habitats

The species occurs at both intertidal and inland sites. Reported soil total salts in ‰ or g/kg are mixed-soil measures and remain separate from NaCl nutrient-solution concentrations; the review’s <5, 15–20, and >20 g/kg values remain secondary field summaries. [S01, S02, S07, S09, S11]

Salinity experiments

Guo et al. (2020) grew wild Dongying seeds in sand pots from sowing to mature seed under 200 mM pure NaCl nutrient solution, raised gradually and then supplied twice daily at three times the pot water-holding capacity. At 103–110 days, the study measured reproductive plants, ovule starch and sugars, and photosynthetic variables; treated plants produced larger seeds. Pore-water concentration was not measured directly. [S04]

Guo et al. (2019) separately grew Dongying-source plants in sand pots from sowing with 200 mM pure NaCl nutrient solution. Vegetative growth was assessed at day 100, followed by observations of flowers and seeds. At day 100, shoot dry mass was 40.7% above the control; fresh mass, height, and primary-branch number were 67.6%, 13.3%, and 12.3% greater, respectively. The study also recorded peak flowering at day 145 and later seed traits. The dry-mass comparison supports actual biomass gain rather than survival judged only at harvest. This protocol is a sand-pot result, not a field root-zone concentration. [S05]

Song et al. (2009) pre-cultured seedlings from Yellow River Delta intertidal and inland populations for 55 days, then gradually applied 10–600 mM NaCl for 18 days. Table 2 shows positive shoot and root relative growth rates at 600 mM in both populations, although rates were lower than at 100–200 mM. Intertidal root dry mass was lower at 600 mM, and leaf water content in the inland population declined at 400 and 600 mM. In a separate sand-culture experiment, Song et al. (2011) found lower shoot dry mass after 22 days at 600 mM; that study did not calculate net growth from initial and final dry mass, so this endpoint alone does not establish sustained growth. [S07, S09]

In Mori et al.’s hydroponic study, post-germination seedlings received nutrient solutions containing 0–800 mol m−3 NaCl for 46 days. Total dry mass was greatest at 50–200 mol m−3, declined above 200, and was extremely low at 800 mol m−3. The control used tap water with small amounts of Na and Cl; this paper reports mol m−3, while other studies use mM or soil-salinity measures. [S08]

Guo et al. (2021) compared plants inoculated with Sphingomonas prati or S. zeicaulis against uninoculated plants and reported 0‰ and 20‰ NaCl. At 20‰, dry mass in the S. prati-inoculated group was 102.25% higher than in uninoculated plants; the abstract also lists height, root length, fresh mass, and physiological endpoints. It does not report plant stage, culture medium, the basis for the salt solution, or treatment duration. The contrast includes an inoculation effect and cannot be generalized to uninoculated plants or converted to mM. [S06]

A one-time Yellow River Delta field survey compared green supratidal and purple intertidal ecotypes. Total salts in surface-site soils were 3.84±1.63‰ and 11.92±1.59‰, respectively; the paper also compared above- and belowground growth. These ‰ values describe mixed soil salts at plot scale, without a time series or a fixed exposure measurement for individual roots. They are not pure-NaCl solutions and are not converted to controlled-treatment concentrations. [S03]

Song and Wang’s review cites Gu (1999) for field communities in which S. salsa was not recorded below 5 g/kg dry soil, was described as suitable at 15–20 g/kg, and could dominate above 20 g/kg. The review does not identify the original plots, sampling duration, or analytical method. These total-soil-salt descriptions are secondary ecological information, not growth-treatment results. [S11]

The databases combine 800, 600, 400, and 200 mM entries with ‰ and mg/L values from independent studies. Song et al. (2009) measured positive relative growth at 600 mM, whereas growth was extremely poor at 800 mol m−3 in Mori et al. Database links also include an inoculation experiment or a biomass-processing paper, so the entries do not form a single dose–response curve. [S10, S14]

Song et al. (2008) exposed brown and black dimorphic seeds from intertidal and inland populations to 0.6–35.1 ppt NaCl for 20 days, with a separate 20-day sand-pot test of emergence and seedling growth. Germination of brown seeds exceeded 96% in both populations across the salinity treatments; 35.1 ppt significantly shortened shoots, with a larger effect on inland seedlings. Germination and short-term seedling growth are distinct endpoints. [S01]

Liu et al. (2020) measured root and leaf Cl− in Yellow River Delta intertidal and inland plants and treated seedlings from both populations with 0 or 300 mM NaCl for 7 or 35 days. Total soil salt was 12.35±1.91 g/kg at the intertidal site and 2.50±0.34 g/kg inland; root Cl− in field intertidal plants was about 2.9 times the inland value. Controlled studies also measured root/leaf Cl−, expression of SsCLCc, Ssperoxidase64, and SsCASP, and root endodermal structure. Gene and barrier roles are the authors’ interpretation, not single-factor functional proof. [S02]

Guo et al. (2018) sowed Yellow River Delta seeds in sand and applied 0, 200, or 400 mM NaCl from sowing through seed maturity, with 12 pots per treatment. The study measured flowering, seed set/yield, and ions in reproductive tissues; reproductive performance was greatest at 200 mM, with smaller gains at 400 mM. This full-cycle experiment differs in material and endpoints from the vegetative-growth trial of Guo et al. (2019). [S18]

Reported mechanisms and ecological adaptation

Studies report ovule and seed development, ion accumulation in leaves and roots, Cl− transport, and expression of genes related to root barriers. Populations differed in root/leaf Cl− allocation; the roles proposed for gene expression and the Casparian band remain interpretations that require functional tests. [S01, S02, S03, S04, S05, S06, S07, S08, S09]

Conclusions and evidence gaps

Sources support natural intertidal and inland populations. The 200 mM pure-NaCl sand-pot study continues into reproduction and covers the broadest developmental span among the available sustained-growth records. Short seedling experiments reached 600 mM, while Mori et al. reported very poor growth at 800 mol m−3; each result applies to its study’s stage and duration. Redondo-Gómez et al. (2008) applied 900 mol m−3 NaCl in S. splendens seed-germination tests, followed by a seedling-growth experiment with a maximum of 400 mol m−3. That paper studied S. splendens and provides no results for S. salsa. [S01, S02, S03, S04, S05, S07, S08, S09, S19]

Salinity research · Evidence records

11

Materials, original units, exposure durations and limitations are retained. These records do not form a single tolerance threshold.

M1

Salinity-related evidence

[S04]
Material
Wild S. salsa seed from Dongying, Shandong; annual plants grown from sowing through mature seed formation.
Salinity metric
Pure NaCl added in nutrient solution
Treatment / value and unit
200 mM NaCl
Medium and exposure
Sand culture; salinity was gradually increased and saline nutrient solution was applied twice daily at three times the pot water-holding capacity to maintain relatively stable sand salinity.
Life stage
Germination through vegetative growth and mature seed production
Duration
From sowing through mature seed formation; reproductive measurements reported around 103–110 days after sowing
Measured endpoint
Whole-plant growth and reproduction, seed number/size, leaf/flower/ovule photosynthesis and carbohydrate measures
Result
Treated plants produced larger seeds than controls. The 2020 paper cites Guo et al. (2018) for earlier increases in seed number, seed weight, and mean individual seed mass. The 2020 experiment itself measured reproductive-stage plant and ovule starch and sugars and leaf photosynthesis at 200 mM.
Scope and limitations
The full-cycle treatment supports growth and reproduction at the tested nominal dose. Salt was increased gradually and pore-water concentration in the sand was not independently measured; this is not a field threshold.
M2

NaCl sand-pot growth and reproductive experiment

[S05]
Material
S. salsa from Dongying; plants grown in sand.
Salinity metric
Pure NaCl added in nutrient solution
Treatment / value and unit
200 mM NaCl
Medium and exposure
Sand pots with nutrient-solution treatment; control and 200 mM groups began at sowing, and saline solution was applied continuously according to the study protocol.
Life stage
Vegetative growth through flowering and seed development
Duration
Biomass at 100 days after sowing; flowering measured at 145 days
Measured endpoint
At 100 days after sowing, whole-plant shoot fresh/dry mass, height, and primary branches; later flowers, pollen, and mature seeds.
Result
At 100 days after sowing, the 200 mM treatment did not suppress growth: shoot dry mass was 40.7% above control; fresh mass, height, and primary branches were 67.6%, 13.3%, and 12.3% greater. The full study also recorded flowering, fruiting, and seed traits.
Scope and limitations
Dry-mass gain supports real growth rather than residual endpoint mass alone. Sand-pot results are not a field root-zone threshold.
M3

Short-term seedling NaCl and waterlogging experiment

[S09]
Material
Seedlings from intertidal and inland saline populations of S. salsa.
Salinity metric
Pure NaCl concentration
Treatment / value and unit
1, 200, and 600 mM NaCl
Medium and exposure
Sand culture with drained or waterlogged conditions.
Life stage
Seedlings
Duration
22 days
Measured endpoint
Seedling response and ion distribution; abstract does not provide net growth at 600 mM
Result
600 mM was the highest treatment, but the accessible abstract does not establish sustained net growth at that concentration.
Scope and limitations
The abstract reports NaCl concentrations and treatment duration but gives no sustained net-growth data above 200 mM; maximum exposure or survival alone does not demonstrate sustained growth at higher concentrations.
M4

Short-term seedling NaCl growth experiment in two populations

[S07]
Material
S. salsa seedlings from Yellow River Delta intertidal and inland saline-soil populations, pre-cultured for 55 days
Salinity metric
Pure NaCl concentration
Treatment / value and unit
10, 100, 200, 400, and 600 mM NaCl
Medium and exposure
Greenhouse sand culture with NaCl added gradually; nutrient solution was replenished weekly
Life stage
Seedlings
Duration
18 days of salinity treatment
Measured endpoint
Shoot and root dry mass, relative growth rate, leaf water content, branching, and ion accumulation
Result
Both populations retained positive shoot and root relative growth rates at 600 mM, although rates were lower than at 100–200 mM. Intertidal root dry mass at 600 mM was lower than at 10 mM; leaf water content in the inland population declined at 400 and 600 mM.
Scope and limitations
Short-term greenhouse sand culture of seedlings; starting biomass and responses differed by population. The 600 mM result is an 18-day relative growth rate, not evidence of long-term full-cycle tolerance or a field root-zone threshold.
M5

Hydroponic NaCl growth-curve experiment

[S08]
Material
The hydroponic experiment used post-germination seedlings; population source was not reported.
Salinity metric
Pure NaCl in hydroponic nutrient solution
Treatment / value and unit
2–800 mol m−3 NaCl in hydroponic nutrient solution, as reported; highest growth at 50–200 mol m−3, declining above 200 mol m−3
Medium and exposure
Hydroponic nutrient solution with graded NaCl.
Life stage
Vegetative plant growth
Duration
Each of eight NaCl treatments lasted 46 days; concentrations above 100 mol m−3 were raised gradually every two days. The experiment was duplicated, with six seedlings transplanted per container.
Measured endpoint
Plant growth and dry matter
Result
The abstract reports increasing growth from 2 to 200 mol m−3, greatest growth from 50 to 200, and decline above 200; growth was very poor at 800 mol m−3.
Scope and limitations
Nutrient solution was prepared with tap water, which contained small amounts of Na and Cl; results apply to the tested hydroponic material and protocol. Total dry weight was extremely low at 800 mol m−3, so this treatment does not indicate positive sustained growth.
M6

Field habitat soil-salt measurements

[S03]
Material
Wild green supratidal and purple intertidal S. salsa ecotypes from the Yellow River Delta
Salinity metric
Total salts in mixed field soil; original unit retained
Treatment / value and unit
Supratidal: 3.84±1.63‰; intertidal: 11.92±1.59‰ (soil total salts, as reported; not a solution concentration).
Medium and exposure
Surface-site soils from the supratidal and intertidal zones of the Yellow River Delta, as reported in Table 2
Life stage
Natural field populations
Duration
One survey; no time series reported
Measured endpoint
Site soil total-salt concentration and plant growth comparison
Result
Total soil salts were 11.92±1.59‰ at intertidal sites and 3.84±1.63‰ above the tide line.
Scope and limitations
Mixed-soil metric at plot scale, not NaCl solution or an individual-root threshold; no conversion to mM.
M7

Endophyte-inoculation growth study, abstract-level

[S06]
Material
S. salsa plants inoculated with Sphingomonas prati or S. zeicaulis.
Salinity metric
NaCl salinity reported in per mille (‰), basis not specified in abstract
Treatment / value and unit
0‰ and 20‰ NaCl, as stated in the abstract
Medium and exposure
Plants with and without endophyte inoculation were compared; the abstract provides no culture medium or salt-solution preparation method.
Life stage
Plant growth; stage not specified
Duration
Not reported
Measured endpoint
Height, root length, fresh/dry mass, and physiological traits
Result
At 20‰, dry mass in the S. prati-inoculated group was 102.25% higher than in uninoculated plants; other growth measures were also reported.
Scope and limitations
The full text was restricted, the basis of ‰ and duration are unknown, and an inoculation response is not generalized to uninoculated plants or converted to mM.
M8

Secondary review of field soil salinity

[S11]
Material
S. salsa field-community reports cited secondarily by Song and Wang (2015), attributed to Gu (1999).
Salinity metric
Total salts per dry-soil mass
Treatment / value and unit
<5 g/kg (not observed); 15–20 g/kg (reported suitable range); >20 g/kg (reported dominance)
Medium and exposure
Song and Wang’s (2015) review cites Gu (1999) for natural saline-soil communities; the original plots and sampling methods are not reported.
Life stage
Natural populations/community occurrence
Duration
Secondary description of long-term habitat; the original sampling duration is not reported
Measured endpoint
Occurrence and reported habitat suitability or dominance
Result
Song and Wang’s review cites Gu (1999) as reporting no S. salsa below 5 g/kg dry soil, a stated suitable range of 15–20 g/kg, and potential dominance above 20 g/kg.
Scope and limitations
The review does not specify the original plots, sampling duration, or analytical method. Total soil salts cannot be converted to NaCl solution concentration or used to infer plant growth in controlled experiments.
M9

Salt treatment of germination, emergence, and early growth

[S01]
Material
Brown and black dimorphic S. salsa seeds from intertidal and inland populations in the Yellow River Delta
Salinity metric
NaCl; treatment concentrations reported in ppt and retained as given
Treatment / value and unit
0.6, 5.8, 11.7, 23.4, and 35.1 ppt NaCl
Medium and exposure
Germination: filter paper/nutrient solution in continuous darkness; emergence and seedlings: saline river-sand pots
Life stage
Seed germination, emergence, and 20-day-old seedlings
Duration
20 days for both germination and emergence/growth tests
Measured endpoint
Germination and emergence percentages, shoot length, shoot dry weight, and water/ion measures
Result
Twenty-day germination of brown seeds exceeded 96% in both populations at all salinities; black seeds germinated less often. At 35.1 ppt, shoot length declined substantially, with a larger reduction in inland seedlings; shoot dry weight of brown-seed seedlings was not adversely affected up to 23.4 ppt.
Scope and limitations
Seed color and maternal habitat affected responses. The paper’s NaCl ppt values are not converted to mM; these tests do not represent sustained growth of established plants.
M10

Field root/leaf Cl− allocation and controlled salt treatment

[S02]
Material
S. salsa plants/seedlings from intertidal and inland populations in the Yellow River Delta
Salinity metric
Field soil total soluble salt and Cl− (water extracts); controlled treatment as NaCl concentration
Treatment / value and unit
Field total salts: 12.35±1.91 and 2.50±0.34 g/kg dry soil; treatments: 0 or 300 mM NaCl
Medium and exposure
Field soil and root/leaf Cl− measurements; controlled seedlings received stepwise salt in river-sand pots
Life stage
Wild plants and established seedlings
Duration
Controlled treatments for 7 or 35 days; single field sampling
Measured endpoint
Root/leaf Cl−, related gene expression, and root endodermal structure
Result
Root Cl− in field intertidal plants was about 2.9 times the inland value. Under controlled conditions, 300 mM NaCl increased root and leaf Cl− and altered expression of barrier-related genes and root tissue observations.
Scope and limitations
Field sites are not paired to constant individual root-zone exposures. Gene expression and Casparian-band observations support candidate mechanisms but do not establish gene-level causality.
M11

Full reproductive-cycle NaCl treatment

[S18]
Material
S. salsa seeds and greenhouse plants from the Yellow River Delta
Salinity metric
NaCl concentration in nutrient solution
Treatment / value and unit
0, 200, and 400 mM NaCl
Medium and exposure
Sand culture; NaCl treatment began at sowing and continued to seed maturity
Life stage
Germination, vegetative growth, flowering, and seed maturation
Duration
From April to seed maturity, approximately one growth cycle
Measured endpoint
Flower number, seed set, seed number/yield per plant, seed mass, and reproductive-tissue ions
Result
Reproductive performance was greatest at 200 mM: flower number, seed number, and seed yield increased and flower abortion declined; gains were smaller at 400 mM.
Scope and limitations
One Yellow River Delta seed source, greenhouse natural light, and sand culture; results do not establish a field root-zone threshold.

Linked database records

Database indexing is presented separately from original research evidence.

eHALOPH species database 29141

eHALOPH groups 800, 200, 600, and 340 mM, 0.5%, and 35,100 mg/L under maximum salinity and lists 200 mM and mg/L optimum values; these fields aggregate different units and sources. Mori et al. (2006) found greatest growth at 50–200 mol m−3, decline above 200, and very low growth at 800 mol m−3. The 0 and 20‰ values in Guo et al. (2021) do not support a conversion to 340 mM, and Tan et al. (2023) studied harvested biomass conversion rather than salt treatment of living plants. The 35,100 mg/L value is not linked to an identifiable primary measurement; these database values do not form one dose–response curve. [S14]

China Halophyte Catalogue CNHP_0975; CNHP_0976

The catalogue records coastal saline flats and waterside saline sand and lists 800 mM in its maximum-salinity field. Its habitat wording agrees with natural saline-site studies; the value is a secondary compilation without identifiable independent material, medium, duration, or growth endpoint. [S10]

BioSaline database species page id=126914

Labels S. salsa a true succulent halophyte and coastal plant; records saline soils, beach/lakeshore habitat, annual habit, morphology, phenology, and uses. It supplies qualitative habitat and classification context, not a salt concentration, and does not replace primary habitat or treatment evidence. [S15]

Shandong Normal University halophyte resource database GRHC_H_1919; GRHC_H_1847; GRHC_H_1697; GRHC_H_1685; GRHC_E_1406

Five exact-name pages record saline-alkaline soil, sandy beaches, and the raw provider soil-salt field “0.4-0.6%”. The descriptions are repeated within one database source chain; denominator and measurement method are unspecified, so the percentage is not converted to a solution concentration or treated as an independent experiment. [S16]

FloraVeg.EU baseline_row_2321

FloraVeg has a species entry, with a European EUNIS saline or brackish relation count of 0; the record supplies no saline-habitat association for the species. [S17]

Preslia regional inventory supplement row 40

The regional inventory lists category Ia, halophyte_evidence=true, and regional salinity indicator S=7. These are regional ecological indicators, not an additional independent field population or a salt concentration; they cannot be converted to NaCl concentration or a transferable plant response. [S12]

Literature & sources

19

Source IDs are local to this species and content version.

S01Primary field study

Song, J., H. Fan, Y.Y. Zhao, Y.H. Jia, X.H. Du & B.S. Wang (2008). Effect of salinity on germination, seedling emergence, seedling growth and ion accumulation of a euhalophyte Suaeda salsa in an intertidal zone and on saline inland. Aquatic Botany 88(4):331–337.

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S02Primary field study

Liu et al. (2020). Chloride allocation in the euhalophyte Suaeda salsa from different habitats in field and controlled saline conditions. Aquatic Botany 167:103292.

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S03Primary field study

Song, Y., J. Liu, J. Wang & F. Liu (2021). Growth, Stoichiometry, and Palatability of Suaeda salsa From Different Habitats Are Demonstrated by Differentially Expressed Proteins and Their Enriched Pathways. Frontiers in Plant Science 12:733882.

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S04Primary experimental study

Guo, J., M. Du, C. Lu & B. Wang (2020). NaCl improves reproduction by enhancing starch accumulation in the ovules of the euhalophyte Suaeda salsa. BMC Plant Biology 20:262.

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S05Primary experimental study

Guo, J., X. Dong, G. Han & B. Wang (2019). Salt-enhanced reproductive development of Suaeda salsa L. coincided with ion transporter gene upregulation in flowers and increased pollen K+ content. Frontiers in Plant Science 10:333.

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S06Primary experimental study

Guo, J., Y. Chen, P. Lu, M. Liu, P. Sun & Z. Zhang (2021). Roles of endophytic bacteria in Suaeda salsa grown in coastal wetlands: Plant growth characteristics and salt tolerance mechanisms. Environmental Pollution 287:117641.

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S07Primary experimental study

Song, J., M. Chen, G. Feng, Y.H. Jia, B.S. Wang & F.S. Zhang (2009). Effect of salinity on growth, ion accumulation and the roles of ions in osmotic adjustment of two populations of Suaeda salsa. Plant and Soil 314(1–2):133–141.

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S10Database

China halophyte catalog 2025, Suaeda salsa entry (duplicate route in frozen input).

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S11Review

Song, J. & B. Wang (2015). Using euhalophytes to understand salt tolerance and to develop saline agriculture: Suaeda salsa as a promising model. Annals of Botany 115(3):541–553.

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S13authoritative_flora

Flora of China Editorial Committee. 2003. Suaeda salsa (L.) Pall. Flora of China 5:389–394.

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