Review Reports - Seed Germination as an Adaptive Response in Halophytes

Round 1

Reviewer 1 Report (Previous Reviewer 3)

Comments and Suggestions for Authors

Review: Seed Germination as an Adaptive Response in Halophytes by Keriman Şekerci, Nahoko Higashitani, Atsushi Higashitani, and Ismail Turkan present their perspective on the characteristics of halophyte seed germination in relation to salinity.
The manuscript contains the necessary sections and is formatted according to the rules, but the authors omitted some valuable information, making this review incomplete. Furthermore, the image used in the review presents a one-sided view and does not address this fascinating aspect of plant adaptation, but rather germination in general.
To address this, the review needs to be expanded in several areas:
1. An evolutionary approach. The causes and cases of this adaptation, taking into account the characteristics of the species and origin, and the mode of adaptation, represent at least two strategies—accumulator plants and excluder plants. This is not reflected in any way, and examples and diversity are not provided. There is no analysis of life forms or a discussion of the relationships and differences between succulents and halophytes. 2. Botanical approach (partially reduced to an extremely weak descriptive form without careful analysis). It is necessary to cite families and species with resistance, and describe halophytic forms of other organisms, such as bacteria and fungi, that aid survival in harsh conditions.
3. Physiological approach - analyze and illustrate turgor regulation, provision of free water, the presence of specialized vacuoles and molecular systems that ensure resistance.
4. The structural features of halophyte seeds are well studied and should be presented and discussed.
5. Groups with different strategies should be cited.
6. The discussion of Arabidopsis in an article on halophytes seems odd - you could look at its closest relatives, such as Thyrengiella.
7. The germination strategies of licorice and sea barley, or another species germinating in salt, should be compared. This area is well-studied, and an incomplete presentation discredits and misleads, suggesting a lack of knowledge in this field, or at least its limitations.
Recently, there have been no reviews from this perspective, and therefore, if the authors can do this work properly, it could be useful, not in its current form, but with careful development. I believe these authors have the experience to contribute these important elements, which will increase interest in this area and support their personal reputation and the journal's standing.

Author Response

We sincerely thank the reviewer for their thorough and constructive evaluation of our

manuscript. We appreciate the insightful suggestions, which have significantly improved

the quality, depth, and scope of our review. Below, we address each point in detail.

Comment 1.

An evolutionary approach. The causes and cases of this adaptation, taking into account the

characteristics of the species and origin, and the mode of adaptation, represent at least two

strategies—accumulator plants and excluder plants. This is not reflected in any way, and

examples and diversity are not provided. There is no analysis of life forms or a discussion of

the relationships and differences between succulents and halophytes.

Response:

We thank the reviewer for this insightful suggestion. We have now expanded the manuscript

to include a dedicated section on the evolutionary strategies of halophytes, specifically

distinguishing between salt "accumulators" and "excluders." Table 1 has been added to

categorize the mentioned species according to these evolutionary strategies.

Comment 2.

Botanical approach (partially reduced to an extremely weak descriptive form without careful

analysis). It is necessary to cite families and species with resistance, and describe

halophytic forms of other organisms, such as bacteria and fungi, that aid survival in harsh

conditions.

Response:

We appreciate this important point. The manuscript has been revised to include:

• Expanded taxonomic coverage, with explicit mention of:

o Major halophytic families (e.g., Amaranthaceae, Brassicaceae, Poaceae) in

Table 1 and Table 2

• A new dedicated subsection on seed-associated microbiomes, including line 173-

200:

o Endophytic bacteria

o Halotolerant PGPR (e.g., Bacillus, Pseudomonas)

o Arbuscular mycorrhizal fungi (AMF)

Comment 3.

Physiological approach - analyze and illustrate turgor regulation, provision of free water, the

presence of specialized vacuoles and molecular systems that ensure resistance.

Response:

We fully agree and have substantially expanded this section:

• Added detailed discussion in Figure 2 and line 257-272:

o Turgor regulation under saline conditions

o Biophysical states of water (bound vs free water)

• Expanded molecular perspective to include:

o ABA-mediated regulation

o ROS detoxification systems

o Protective molecules (e.g., LEA proteins, antioxidant enzymes)

These revisions provide a more mechanistic and integrative physiological framework.

Comment 4.

The structural features of halophyte seeds are well studied and should be presented and

discussed.

Response:

We agree that structural adaptations are vital to halophyte success. We have significantly

expanded the manuscript to include a detailed discussion of these features. Specifically:

• Table 1 has been updated to include a column on "Structural and Germination

Characteristics," detailing traits such as seed coat (testa) thickness, mucilage

production (myxospermy), and embryo types.

Comment 5. Groups with different strategies should be cited.

Response:

We have revised the manuscript to explicitly cite and categorize halophytes into distinct

groups based on their adaptive strategies. Following the reviewer's suggestion, we now

distinguish between Salt Accumulators (Euhalophytes), Salt Excluders, and Salt Secretors

(Recretohalophytes). We have also expanded our citations to include diverse families such

as Amaranthaceae, Poaceae, and Primulaceae, providing a botanical analysis of how these

groups differ in their life forms (succulents vs. non-succulents) and their specific

mechanisms of resistance. These classifications have been integrated into the text and

summarized in the updated Table 1.

Comment 6.

The discussion of Arabidopsis in an article on halophytes seems odd - you could look at its

closest relatives, such as Thyrengiella.

Response:

We appreciate this observation. We have revised the manuscript to:

• Reduce emphasis on Arabidopsis thaliana remove some sentence. For example“In

contrast, the glycophyte A. thaliana exhibits limited or no germination recovery

following salt stress” line 394-395.

• Replace or complement with halophytic relatives, particularly:

o Eutrema salsugineum

o Schrenkiella parvula

Arabidopsis is now used only where necessary as a genetic reference model, with clear

justification.

Comment 7.

The germination strategies of licorice and sea barley, or another species germinating in salt,

should be compared. This area is well-studied, and an incomplete presentation discredits

and misleads, suggesting a lack of knowledge in this field, or at least its limitations.

Recently, there have been no reviews from this perspective, and therefore, if the authors can

do this work properly, it could be useful, not in its current form, but with careful

development. I believe these authors have the experience to contribute these important

elements, which will increase interest in this area and support their personal reputation and

the journal's standing.

Response:

We have incorporated comparative case studies, including:

• Sea barley (Hordeum marinum)

• Additional halophyte species with contrasting germination strategies

This section now highlights line 281-284, line 287-292

• Species-specific differences

• Ecological trade-offs

• Recovery vs immediate germination strategies

Author Response File: Author Response.pdf

Reviewer 2 Report (New Reviewer)

Comments and Suggestions for Authors

The authors provide a comprehensive summary of the adaptive response and studies done so far to address salt stress in halophytes. However, there are some questions i have for the authors:

Can you provide some more stress not just the NaCl stressed and how halophytes adapt to it. Is the mechanism similar across different salt stress or there is some kinds of convergence.
also, authors should provide a table listing all the previous studies that have been done on different kinds of salt stress and what kind of stress was used.
The long-term impacts of salinity stress on halophyte seed viability and germination were also not discussed which would be useful, especially under prolonged exposure to extreme conditions.
also if authors could discuss more about the interplay between salt stress and temperature stress or any other stress.
some information can be presented in a form of a cartoon specially some of the stress pathways. That would be useful.

Comments on the Quality of English Language

Can be improved.

Author Response

We thank the reviewer for recognizing the value of our review and for the constructive

suggestions.

Comment 1: Can you provide some more stress not just the NaCl stressed and how halophytes adapt

to it. Is the mechanism similar across different salt stress or there is some kinds of convergence.

Response 1:We have expanded the manuscript to include in Table 2:

• Additional salt types:

o Na₂SO₄

o Na₂CO₃ (alkaline stress)

• Discussion on:

o Osmotic vs ionic vs alkaline stress differences

o Evidence of convergent physiological responses (ABA signaling, osmotic

adjustment)

o Stress-specific distinctions (e.g., membrane damage under alkaline stress

Comment 2:also, authors should provide a table listing all the previous studies that have

been done on different kinds of salt stress and what kind of stress was used.

Response 2: A new table (Table 1 and 2) has been added, summarizing:

• Species

• Type of salt stress

• Concentrations used

• Germination responses

This improves clarity and accessibility of existing literature.

Comment 3:

The long-term impacts of salinity stress on halophyte seed viability and germination were

also not discussed which would be useful, especially under prolonged exposure to

extreme conditions.

Response 3: We have added a new discussion on:

• Long-term salinity exposure in line 338-339, line 431-432

Comment 4:

also if authors could discuss more about the interplay between salt stress and

temperature stress or any other stress.

Response 4: We expanded the section on multiple stress interactions, including line 274-

292:

• Salinity × temperature

• Salinity × osmotic stress

• Salinity × waterlogging

We highlight:

• Synergistic vs antagonistic effects

• Ecological relevance in natural habitats

Comment 5: some information can be presented in a form of a cartoon specially some of

the stress pathways. That would be useful.

Response 5: We agree and have in Figure 2:

• Revised existing figures to better represent:

o Stress-response pathways

o Germination regulation under salinity

• Added schematic illustration of:

o Hormonal regulation

o ROS signaling

o Quiescence vs germination transition

Author Response File: Author Response.pdf

Round 2

Reviewer 1 Report (Previous Reviewer 3)

Comments and Suggestions for Authors

Review: Seed Germination as an Adaptive Response in Halophytes
by Keriman Şekerci, Nahoko Higashitani, Atsushi Higashitani, and Ismail Turkan made some edits to the review,
but unfortunately I cannot consider them definitive.
This is because the authors use erroneous concepts that are not only incorrect but also potentially misleading.
The authors should understand that in living systems, there are very few examples of reversibility (one of the few is plasmolysis, cited by the authors, which can only occur in cells with a certain type of cell wall and is reversible in a small number of cases; in most cases, due to damage to the cytoskeleton, it will lead to cell death). Therefore, if a seed begins to germinate and is exposed to high salinity conditions, it will die. However, many halophytes are characterized by a different situation: the seed can swell, which is not the same as germinating. If exposed to drought or salinity, it will not germinate, but will retain the ability to germinate when exposed to favorable conditions.
The article contains many errors based either on misunderstandings or translation problems.
Particularly shocking is the use of such an oxymuron as "extremophile seeds." Halophytes do not have special seeds; many non-halophyte plants can withstand unique adverse conditions, but not all halophytes can. Please review the text carefully.
My request to clarify the families was fulfilled, but apparently without a serious understanding of the differences in the physiology of different seeds. Okay.
Let's examine the differences.
Dicotyledonous seeds, as is clear, are not all the same. For example, some lack endosperm in the mature state, which affects swelling and subsequently germination. Let's consider true seeds (not fruits, as in cereals).
Seeds contain storage substances, namely starch, lipids in the form of lipid bodies, and proteins in the form of protein bodies. The ratio of these storage substances varies dramatically among different species.
For example, oilseeds store very poorly because the oil oxidizes and becomes toxic. Clearly, swelling of such seeds is critical for survival.
Seeds with a high starch content store better and can withstand prolonged and multiple variations in swelling.
Seeds, which primarily contain proteins, can oxidize and also hinder embryo development.
This is only one aspect, and it isn't even reflected in the general context. The stages of swelling are associated with changes in water availability (in seeds, it exists in at least three states), and this is directly related to physiology.
Reflect these aspects in your work, and edit the drawings to reflect these terms.
Expand the concept to include how exactly halophyte seeds differ—this remains unclear. The work remains raw and, due to the importance of the topic being discussed, still requires the attention of the authors.

Author Response

We sincerely thank the reviewer for their thorough and constructive evaluation of our

manuscript. We appreciate the insightful suggestions, which have significantly improved

the quality, depth, and scope of our review. Below, we address each point in detail.

Reviwer 1

Comment 1

This is because the authors use erroneous concepts that are not only incorrect but also potentially misleading.
The authors should understand that in living systems, there are very few examples of reversibility (one of the few is plasmolysis, cited by the authors, which can only occur in cells with a certain type of cell wall and is reversible in a small number of cases; in most cases, due to damage to the cytoskeleton, it will lead to cell death). Therefore, if a seed begins to germinate and is exposed to high salinity conditions, it will die. However, many halophytes are characterized by a different situation: the seed can swell, which is not the same as germinating. If exposed to drought or salinity, it will not germinate, but will retain the ability to germinate when exposed to favorable conditions.

Response 1

Thank you for your clarification. We have replaced the term 'reversible germination' with 'germination recovery' or 'quiescence' throughout the text."

Line 25, 71, 76, 276-277, 364, 396, 403, 528

Comment 2

The article contains many errors based either on misunderstandings or translation problems.
Particularly shocking is the use of such an oxymuron as "extremophile seeds." Halophytes do not have special seeds; many non-halophyte plants can withstand unique adverse conditions, but not all halophytes can. Please review the text carefully.

Response 2

We appreciate the reviewer’s correction regarding the terminology. We agree that 'extremophile seeds' is an inaccurate term.In the revised manuscript, we have removed this term and replaced it with more precise language, such as “halophyte seeds”.

Line 27.

Comment 3

My request to clarify the families was fulfilled, but apparently without a serious understanding of the differences in the physiology of different seeds. Okay.
Let's examine the differences.
Dicotyledonous seeds, as is clear, are not all the same. For example, some lack endosperm in the mature state, which affects swelling and subsequently germination. Let's consider true seeds (not fruits, as in cereals).
Seeds contain storage substances, namely starch, lipids in the form of lipid bodies, and proteins in the form of protein bodies. The ratio of these storage substances varies dramatically among different species.
For example, oilseeds store very poorly because the oil oxidizes and becomes toxic. Clearly, swelling of such seeds is critical for survival.
Seeds with a high starch content store better and can withstand prolonged and multiple variations in swelling.
Seeds, which primarily contain proteins, can oxidize and also hinder embryo development.
This is only one aspect, and it isn't even reflected in the general context. The stages of swelling are associated with changes in water availability (in seeds, it exists in at least three states), and this is directly related to physiology.
Reflect these aspects in your work, and edit the drawings to reflect these terms.
Expand the concept to include how exactly halophyte seeds differ—this remains unclear. The work remains raw and, due to the importance of the topic being discussed, still requires the attention of the authors.

Response 3

We sincerely thank the reviewer for this profound and technically rigorous critique.

We have revised the manuscript to move beyond a generic germination model. New sections and Figure 3 now detail how storage ratios (starch vs. lipids vs. proteins) dictate seed survival under salinity.

Line 181-188, 296-311, 317-322 and Figure 3.

Author Response File: Author Response.pdf

Round 3

Reviewer 1 Report (Previous Reviewer 3)

Comments and Suggestions for Authors

Review: Seed Germination as an Adaptive Response in Halophytes
The authors, Keriman Şekerci *, Nahoko Higashitani, Atsushi Higashitani, and Ismail Turkan, failed to demonstrate the differences and diversity in the review and its revision.
The added illustration demonstrates a critical misunderstanding of the differences between halophyte seeds, their germination responses, and the reasons for these differences. There is no description or discussion of the added image, but what is shown there makes the situation even worse than the earlier images and materials.
I cannot find any possible reason why the authors failed to study the issue and correct it during this time.
Furthermore, the authors failed to present any differences between halophyte and glycophyte seeds, or between different groups of halophytes.
Meanwhile, this is a fairly well-researched topic, and such errors are unacceptable. I don't have the space to cite numerous classifications and studies, and this clearly won't quickly improve the depth of understanding of the issue under study, so I'll provide a quote with generally known information (HALOPHYTES: RESOURCES, ECOLOGICAL CHARACTERISTICS, AREAS OF USE N.Z. Shamsutdinov):
Halophytes include plants belonging to various life forms that are capable of completing a full life cycle and reproducing in soil solution salinity with an electrical conductivity of 8-10 dS/m (Aronson, 1985).
Halophytes differ in their response to soil salinity—halotolerance. The range of soil solution mineralization within which a given plant can grow normally and have the ability to reproduce varies among species. According to N.I. According to Akzhigitova (1982), halophytes can be divided into the following groups: 1 - hyperhalophytes, 2 - euhalophytes, 3 - hemichalophytes, 4 - haloglycophytes.
She classifies plants from excessively saline soils as hyperhalophytes. Their ontogenesis occurs better in saline conditions. These halophytes function and reproduce normally in very highly saline soils with a dry residue of: 2.3 - 3.0 (3.5); C1>0.23% for chloride-sulfate salinity or, respectively, 1.8 - 2.3%; C1>0.23% for sulfate-chloride salinity.
Characteristic and widespread halophytes of this group include the following species, usually dominant in solonchaks: Halocnemum strobilaceum (Pall.) Bieb., Halostachys caspica C.A. Mey, Kalidium caspicum (L.) Uno. — Sternb., Climacoptera crassa (Bieb.) Botsh., C. lanata (Pall.) Botsh., Suaeda arcuata Bunge, S. salsa (L.) Pall., Salicornia europaea L., Frankenia pulverulenta L., and others.

The euhalophytes group includes plants that successfully grow in saline soils across a wide range of soil solution mineralization. Halophytes of this group are also well adapted to highly saline soils; however, they develop better and dominate in communities with lower substrate salinity, with dry residue 1.8–2.3 (2.5); Cl>0.1–0.23% for chloride-sulfate salinity or 1.3–1.8%; Cl>0.1–0.23% for sulfate-chloride salinity, respectively. Based on their adaptation method, these are primarily salt-accumulating and salt-secreting halophytes.

N. I. Akzhigitova (1982) classified plants of moderately saline soils as belonging to the third group, hemigalophytes. They develop normally and form relatively high phytomass indices at the following soil salinities: dry residue 1.3–1.8 (2.0); C1>0.03–0.1% or, respectively, 0.8–1.3%; C1>0.03–0.1%.
The haloglycophyte group includes plants with low salt tolerance. Haloglycophytes develop normally and regenerate at dry residues of 0.3–0.8 (1.0); C1>0.01–0.03% or, respectively, dry residue 0.1–0.2%; C1>0.01–0.03% (Akzhigitova, 1982).
According to N.I. According to Akzhigitova (1982), halophytes are divided into the following groups based on their response to moisture: halomesophytes, halomesophytes, halomesoxerophytes, and haloxerophytes. Halomesophytes are predominantly plants of saline, humid habitats with groundwater levels close to the soil surface. These include coastal areas of salt lakes and wet salt marshes, where the groundwater table is at a depth of 0.5–1.0–3.0 m.
Halohemimesophytes include plants of less humid habitats, where the groundwater table is at a depth of 1.5–2.0 (2.5) m.
The halophyte group, halomesoxerophytes, exhibits a broader range of halotolerance in humid conditions, developing normally at a groundwater level of 1.5–4.0 m.
Haloxerophytes are a fairly large group of plants that inhabit saline soils, where the groundwater table is sufficiently deep to have no practical impact on the growth, development, and productivity of halophytes.
Halophytes are heterogeneous in their bioecological, physiological, and biochemical properties, as well as their economically useful characteristics. Ecologically distinct hyperhalophytes, euhalophytes, hemigalophytes and haloglycophytes react differently to excess salinity of the soil solution.

I believe that further work on this manuscript will only worsen the situation, and publication of this review will have negative repercussions and could affect the journal's reputation. I consider it necessary to cease work on this manuscript.

Author Response

We would like to extend our deepest gratitude for your rigorous and critical evaluation of our manuscript. We have carefully considered your comments regarding the lack of diversity and ecological depth in our initial version. Your recommendation to incorporate the classic ecological frameworks of N.I. Akzhigitova (1982) and N.Z. Shamsutdinovhas been a turning point for this review.

We have fundamentally restructured the manuscript to bridge these seminal ecological classifications with modern physiological mechanisms. We believe these extensive revisions have transformed the review into a comprehensive and scientifically grounded work that accurately reflects the diversity of halophyte germination strategies.

Below is our point-by-point response to your concerns:

Comment 1: " The added illustration demonstrates a critical misunderstanding of the differences between halophyte seeds, their germination responses, and the reasons for these differences...."

Response: We have completely redesigned Figure 3 to address the reviewer’s concerns regarding the lack of diversity and scientific depth. The previous version provided a generalized view that did not account for the heterogeneous nature of halophytes.

We highlight that while salinity leads to irreversible membrane damage and -amylase inhibition in glycophytes, halophytes possess unique traits such as specialized fatty acid profiles (e.g., maintaining linoleic acid) and seed coat heteromorphism (e.g., dimorphic seeds) to ensure reproductive success in saline environments.

Figure 3, Line 321-327, Line 328-336, Line 342-348,

Comment 2: " Furthermore, the authors failed to present any differences between halophyte and glycophyte seeds, or between different groups of halophytes.

Response 2: We completely agree that a review on halophytes must be anchored in their ecological niches to be scientifically valid. Following your guidance, we have integrated the eco-hydrological and salinity-based classifications into the core of our discussion:

Textual Alignment with Akzhigitova (1982):We have integrated a comprehensive discussion in the introduction part bridging these physical seed mechanisms with the classic groundwater-depth classifications (halomesophytes, halohemimesophytes, halomesoxerophytes, and haloxerophytes). We clearly demonstrate how the functional data presented in legend of Table 1 (Salt Accumulators vs. Excluders) and Table 2 (Obligate vs. Facultative) provide the exact physiological basis for these historical ecological groupings.

Line 45-69, Line 89-91, Line 127, Line 157, Line 190, Line 192, Line 193, Line 201, Line 220, Line 231, Line 244, Line 279, Line 313, Line 326, Line 333, Line 357, Line 490, Line 499, Line 523, Table 1 and 2 legend.

We believe these extensive clarifications remove any perceived "uniformity" in our model and provide the exact ecological and evolutionary depth the reviewer rightfully demanded.

We have taken your warning regarding the journal's reputation very seriously. By incorporating the authoritative ecological classifications you suggested, we believe we have moved beyond a surface-level summary to a high-level academic synthesis. We hope these revisions demonstrate our commitment to scientific accuracy and depth.

Thank you again for your invaluable contribution to the improvement of this work.

Author Response File: Author Response.pdf

Round 4

Reviewer 1 Report (Previous Reviewer 3)

Comments and Suggestions for Authors

The questions are taken into account in the manuscript.

This manuscript is a resubmission of an earlier submission. The following is a list of the peer review reports and author responses from that submission.

Round 1

Reviewer 1 Report

Comments and Suggestions for Authors

In this manuscript, Şekerci et al. present an interesting and timely overview of seed germination in halophytes, highlighting the differences from the same process in glycophytes. High soil salinity generally causes an irreversible inhibition of germination in salt-sensitive plants, whereas in halophytes it often induces a reversible and tightly controlled suppression of germination: the seeds maintain their viability and germinate efficiently once salinity levels are reduced. The review describes and discusses the distinctive characteristics of halophyte seeds and the mechanisms regulating germination, dormancy, and recovery, including morphological and structural traits (such as seed heteromorphism, protective seed coats, and mucilage production), as well as physiological and molecular controls (phytohormones and ROS levels, signalling pathways, and the regulation of gene expression). The comparison between the natural process of germination recovery in halophyte seeds and the biotechnological practice of seed (salt) priming is also of particular interest. The information presented is supported by more than 100 appropriate bibliographic references, half of which, approximately, were published within the last five years.

Seed germination and seedling establishment are crucial stages of the plant life cycle, highly sensitive to abiotic stress, and have therefore been the subject of extensive research, resulting in numerous publications, including high-quality review articles. The same can be said about the mechanisms of plant responses to salt stress. However, relatively few recent reviews have focused specifically on the behaviour of halophyte seeds during germination, dormancy and recovery. The present article thus could make a valuable contribution by helping to fill this gap.

Nevertheless, before the manuscript can be accepted for publication, some minor revisions are required to address formal issues (e.g., the lack of consecutive numbering in the citations in the text, or the absence of italics for some scientific names) and, more importantly, to improve the quality of the English language and style (see "Comments on the Quality of English Language")

Comments for author File: Comments.pdf

Comments on the Quality of English Language

The text should be carefully revised to correct basic grammatical errors (incorrect verb forms, inappropriate word usage, etc.) and to eliminate unclear or nonsense sentences (for example, sentences without a verb or missing some words) that sometimes reduce comprehension of the scientific content. Some examples of such mistakes and unclear sentences are indicated in the attached annotated copy of the manuscript.

Reviewer 2 Report

Comments and Suggestions for Authors

The manuscript by Sekerci et al. presents an interesting focus on seed germination in halophytes driven by an adaptive decision process. The topic is very timely and interesting; however, the MS presents substantial flaws. The most important limit is the seed dormancy concept that is somehow distorted. Seed dormancy is a trait that, as the authors say, prevents germination under short periods of favorable conditions within unfavorable seasons. In this view the germination inhibition during seed imbibition in the presence of salt cannot be assimilated to a favorable condition for germination. The presence of salt is never favorable for germination. Primary dormancy is acquired during seed maturation, so in this case it is maximum at the dispersal and then released when a seed passes a determined series of conditions. So in the case of halophytes that promptly germinate in water and are instead inhibited in saline environments, there is no dormancy. The fact that dormancy-associated homologous genes are regulated during the inhibition of germination in the presence of salt is evidence that the seed uses the same pathways of dormancy but it doesn’t mean that it is the same process. As for the secondary dormancy, it is acquired when a primary dormancy is removed and the favorable conditions are not met. Even for this case, saline germination inhibition cannot fit, as the seed passes from an unfavorable to favorable condition; that’s it. My idea is that the harsh conditions that are experienced by halophytes are so extreme that the seeds living in this habitat do not need to have a complicated decision process to meet the exact period to germinate; they just need to avoid salinity. For this reason I think that the review should be entirely rethought.

Reviewer 3 Report

Comments and Suggestions for Authors

"Review: Seed Germination as an Adaptive Decision Process in Halophytes" by Keriman Sekerci, Nahoko Higashitani, Atsushi Higashitani, and Ismail Turkan examines specific evolutionary mechanisms that contributed to the survival of some halophytes.
The manuscript is organized according to the requirements and contains the necessary sections. However, its logical construction, focusing on analyzing individual adaptation examples without forming a structured, complex picture or logically constructing interdependencies, makes it mosaic-like and incomplete.
The authors' chosen topic is quite complex, and while some details are thoroughly studied, others remain unclear. A review should a priori have a guiding message, some conclusion, and indications of unresolved issues. However, after reading this manuscript, I find no unifying system, as the diversity of the research subjects is not addressed. Halophytes are extremely diverse, belonging to completely different ecological groups, including dicotyledons, monocotyledons, and even aquatic plants, and their survival and dormancy strategies are completely different. Finally, some plants don't have seeds at all—for example, monocots—but we're dealing with fruits. Among monocots, there are many halophytes, and they have different strategies.
I don't see any analysis of the differences between the botanical variations of seeds; the roles of different tissues, such as endosperm, the specific localization and leaching of hormones during germination, and other botanical, ecological, and functional aspects are not reflected.
For example, the authors don't mention that water in seeds exists in four states, and that changing the percentage ratio between bound and free water is key to triggering swelling and division.

Some questions are puzzling; the authors write that hormone metabolism can occur in seeds. These assertions are surprising, since in dormant seeds, metabolism is reduced to a minimum; everything synthesized was formed and stored during ripening before the loss of free water, and active processes are regulated precisely by its presence. In seeds, we are dealing with passive processes; even root growth is not directly related to division. The increase in root size, or growth as we see it, is first swelling, then elongation, and only then division occurs.

All of these aspects are not discussed in the article, nor is the phenomenon of salt sensitivity and tolerance, which can yield surprising examples, such as licorice, which is completely unable to germinate in the presence of salts, examined in detail.
Finally, the authors completely ignored a number of important studies demonstrating the potential for using saline regions as forage lands, for example, Shamsutdinov, N., & Shamsutdinov, Z. (2008). Halophyte utilization for biodiversity and productivity of degraded pasture restoration in arid regions of Central Asia and Russia. In: Biosaline agriculture and high salinity tolerance (pp. 233-240). Basel: Birkhäuser Basel.
Overall, I find the topic quite relevant, but a high-quality revision of the manuscript would require either a change in focus to the role of priming in halophyte cultivation, in this case with the addition of the biophysics and physiology of seeds and fruits, or a more botanical and ecological perspective, in this case focusing on ecotypes, evolutionary adaptation variants, and an analysis of research prospects in this area.
In its current form, it is not possible to recommend the manuscript for publication.

Reviewer 4 Report

Comments and Suggestions for Authors

The review paper summarized the specialty of halophytes from seed traits and germination perspectives, as well as their underlying physiological and molecular mechanism that adapts to high salinity environments, which is valuable and of significance. But the manuscript has some shortcomings that need to be revised.

Germination decision of halophytes includes germination, dormancy and recovery. Generally speaking, I agree the frame diagram of Figure. As Fig. 1 showed, seed traits including seed coat, mucilage and heteromorphism influence germination, dormancy and recovery. The contents should not be under germination, but to be a subtitle separately.
Under “The impact of saline environment on the seed germination”, the authors should talk about the threshold of salinity that halophytes can bear to germinate and salt type sort of thing. Although some halophytes can’t germinate in high salinity, but most of the halophytes can germinate in above 300mM NaCl salt solutions, even 1M or 1.2M.
In the Dormancy section, the authors just talked about primary dormancy. But most halophytes have the ability to enter secondary/conditional dormancy when their seeds exposed to high salinity conditions.
In the recovery of germination section, the authors didn’t summarize the previous study and the points very well. The salt type, salt concentration and exposure duration in salts closely related to recovery of germination. These points were not discussed.
“Understanding the mechanisms underlying the production of seeds with different structural and functional characteristics of some halo phytes growing in conditions that are not suitable for agriculture is vital for food security, the economy, and the preservation of ecological diversity.” Based on this sentence, it seems that the review paper is about seed production of halophytes.
Figure 1 showed much more information than the last paragraph of Introduction. Therefore, the paragraph should expand a bit and briefly describe Figure 1.
Conclusion section should highlight the strategies or specialty that halophytes adopt to adapt to the high salinity environments.
“Recovery of germination of halophyte seeds is a key determinant for their distribution and population persistence.” The statement is not true. At least provide references.
Use full Latin name of species when the authors first mention the species, not in short. For example, salsugineum.
There are some logic problems or grammar errors in some sentences, which are confusing. The whole paper needs to be revised and improved in English. Take the following sentences for example:

“They are also used as forage, oilseed, medicine, and food are considered good phytoremediators for metal-contaminated saline environments, such as coastal regions.” (second paragraph of 1. Introduction)

“…which can delay or prevent germination until unfavorable conditions.” (second paragraph of 1. Introduction)

“…it is plausible to suggest that halophyte seeds are a very effective means of total adaptive strategies of plants occupying diverse harsh environments…”(second paragraph of 1. Introduction)

This suggestion can be using nutrient-rich food sources. (5. The utilization of halophyte…)