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Review
Peer-Review Record

Molecular Advances in Breeding for Durable Resistance against Pests and Diseases in Wheat: Opportunities and Challenges

Agronomy 2023, 13(3), 628; https://doi.org/10.3390/agronomy13030628
by Kun Luo *, Dejia He, Jiao Guo, Guangwei Li, Boliao Li and Xiulin Chen
Reviewer 2: Anonymous
Reviewer 3:
Agronomy 2023, 13(3), 628; https://doi.org/10.3390/agronomy13030628
Submission received: 22 December 2022 / Revised: 18 February 2023 / Accepted: 20 February 2023 / Published: 22 February 2023
(This article belongs to the Special Issue Advances in Plant Genetic Breeding and Molecular Biology)

Round 1

Reviewer 1 Report

 Luo et al. review manuscript entitled “Breeding Wheat for Durable Resistance against Pest and Disease: Opportunities and Challenges.”  To improve the manuscript please follow the following suggestions.

Line 26: please rewrite the sentence.

Line no: 47 ‘Breed’ replace with breeding of

Authors mentioned Fig.1, Fig.2, and Fig.3 does not present in the manuscript.

 

It is better if data presented in table forms.

Author Response

Response to Reviewer 1 Comments

 

Dear Editor and Reviewers,

Thank you very much for your letter and the comments about our paper submitted to your journal. We would also like to thank the reviewers for their careful and constructive reviews. Soon after receiving your comments, the other authors and I carefully revised the manuscript. The changes made according to the comments from the reviewers are detailed below.

 

Reviewer #1: Luo et al. review manuscript entitled “Breeding Wheat for Durable Resistance against Pest and Disease: Opportunities and Challenges.”  To improve the manuscript please follow the following suggestions.

Response 1: Thank you very much for carefully reviewing our manuscript and kindly giving us many suggestions and comments for improving the readability of this manuscript. In the revision, we carefully followed your corrections.

 

Line 26: please rewrite the sentence.

Response 2: In the revision, we rewrote that section and corrected the illogical points in the introduction.

 

Line no: 47 ‘Breed’ replace with breeding of

Response 3: Yes, as suggested, we reworded that phrase as follows: “breeding of”. (Line 80)

 

Authors mentioned Fig. 1, Fig. 2, and Fig. 3 does not present in the manuscript.

Response 4: We apologize for not uploading the three figures in the original submission. To improve the readability and logic of the figures, in this manuscript, we modified the text and the arrows in the figures and added more content to the descriptions in the figure legends following your suggestion and comments.

It is better if data presented in table forms.

Response 5: Thank you very much for your suggestions. The manuscript has focused on putting together the different molecular related approaches that have been implemented in the direction of breeding wheat varieties that are resistant or tolerant to different biotic stresses at the same time where opportunities exist and challenges thereon. Thus, we prepared three figures to illustrate the regular flow charts of the integration of desirable traits into wheat cultivars in wheat breeding programs, prevalent wheat germplasm improvement strategies and the principal stages in genetic engineering in wheat breeding, and a model summarizing the approaches for future development of wheat cultivars with durable improvements. We hypothesize that these three figures could more accurately describe the recent advances and drawbacks related to the measures associated with molecular breeding in common wheat cultivar improvement for the readers. Please allow us to keep these figures.

 

Reviewer 2 Report

Dear authors,

 

Please find some remarks and questions (en green) below:

L27-28 :

 “…production of bread wheat (Triticum aestivum L.) is of paramount importance to mankind, especially in arid and semiarid areas’’

The production of wheat is important in every area and not more in arid conditions… This is the first crop of the world.

 

 

L37-38 :

 “…In addition, returning straw back into fields also potentially increases the rate of occurrence of plant diseases  and insect pests in the subsequent season of crop growth…”

Personally, I think that this accurate information is not relevant in this part of the text, which present the general risk caused by biotic stress.

 

L42-43

            “…irrational application has considerably enhanced environmental issues and pest resurgence”

                Most of the application are not irrational in the whole world…

 

L43-46

“… rapid increase in the consumption of high-quality foods and products derived from wheat grains, which has significantly promoted the development of wheat resistance and biofortification breeding programs.

There is not link between the increase of wheat consumption in some part of the world and the development of resistant cultivars. From the beginning of the wheat breeding, breeders have tried to develop resistant and tolerant varieties.

 

L62

“…Based on these advances, wheat breeders have given much attention to the development of hybrids…”

Be careful at the definition of hybrids in wheat. By the past, ‘Hybrid” was synonymous with “crossing” but at the present time hybrid means Varieties in first generation sold to the farmer…

 

And as said before, breeders has not waited the molecular advances to give attention to the durable resistance to biotic attackers

 

L66-69

“…Although some of these resistant wheat cultivars could adversely affect the performance of fungal diseases and herbivorous insects, there were still some obstacles that severely limited the large-scale practical  applications of resistant cultivars in agriculture.”

 

Are you talking about resistant cultivars obtained by new molecular techniques or by conventional breeding?

L76

“…by contrasting new… “

Appropriate term?

L84-85

                “….conventional plant breeding programs have always relied on phenotypic characteristics for the development of new cultivars or the improvement of existing cultivars…”

                Conventional breeding of wheat has always relied on phenotypic characteristics,  yes but also on quality of grains and flours analysis…

L86-88

            “….breeders usually spend more time selecting wheat cultivars with the desired traits, while most of the preferred traits are controlled by quantitative trait loci (QTL)”

 

                I doubt that the “most preferred traits are the one of QTL instead of the one of the breeder.

 

L88-90

“…Therefore, against the new background of wheat breeding, conventional wheat breeding cannot fulfill the demands of sustainable agriculture.”

 

I strongly disagree with this statement! Please inform yourself on new varieties obtained all around the world by conventional breeder.

 

L92-93

“…could not only disrupt reproductive isolation”

 

This sentence is not clear

 

L99-100

                “…Thus, large-scale cultivation of GM wheat plants is considered to be a reliable and environmentally friendly approach to combat crop diseases and herbivorous insects in agricultural production.”

“Considered” by you, author? undoubtedly but what about the opinion of the rest of the world?

 

L125-128

“…Although there were some efforts focused on the development of transgenic wheat resistant to different cereal viral diseases  until the early 2000s, the accumulating experimental evidence revealed that it is not a trivial task [23], probably because most viruses are transmitted by herbivorous insects”

 

             Do you mean that this work is trivial because of the aphids? Or the aphid transmission should be considered as an opportunity to stop the epidemic?

 

L151-L155

“…..Meanwhile, a diverse set of foreign genes associated with the qualified traits of abiotic stress tolerance and yield and quality were integrated into elite wheat cultivars with ATMT and callus bombardment techniques, and the cultivars carrying these foreign genes exhibited high yield and quality and exhibited tolerance to different abiotic stresses.”

 

Could you give some example of elite cultivars obtained by these methods?

 

L187

« DDuring »

 

Typing error

 

L216-L217

 

“…wheat is a natural hybrid of Triticum urartu (L.), T. turgidum (L.) 216 and Aegilops tauschii (Coss.)”

Wheat is a natural hybrid of Triticum Urartu (AA), a close parent of Aegilops speltoides (BB)) and Aegilops tauschii (DD). T. turgidum (AABB) is the product of the hybridation between the genomes AA and BB.

 

L217-L219

 

“…it has the largest genome, probably because it has two copies of three groups of 42 chromosomes (2n = 6x = 42, genome formula AABBDD) obtained from each of its ancestors”

 

Each ancestors has not 42 chromosomes but only 2 x 7chr. = 14 chr. Wheat has (3 x 14chr = 42chr.)

 

L231-233

 

“….QRp.slu.4BL, which conferred antibiosis resistance against the bird cherry-oat aphid (Rhopalosiphum padi L.), while QRp.slu.5AL and QRp.slu.5BL conferred tolerance to R. padi

 

? then all the 3 conferred antibiosis resistance to R. padi … why the author use “while”?

 

L280-281

 

“…Taken together, these molecular markers involved in MAS would enable a reliable and feasible approach to detect resistant germplasms and track resistance genes in wheat breeding programs.

 

Finding gene mutation doesn’t mean finding resistance. Most of the mutation have not the expected effect on the phenotype in the field. The most “reliable” mean to detect resistance is to observe the plant in their environment when they face the (a)biotic stress…

 

L298-302

“Moreover, common (hexaploid) wheat is a natural hybrid of Triticum urartu (L.), 298 T. turgidum (L.) and Aegilops tauschii (Coss.), and it has an extremely large genome size 299 (16,000 Mbp), probably because it has two copies of three groups of 42 chromosomes (2n 300 = 6x = 42, genome formula AABBDD) obtained from each of its ancestors and encodes an 301 estimated 164,000 to 334,000 genes [38].”

 

This sentence is the exact repetition of the line L216-L219…

 

L369-371

 

                In comparison with disease resistance gene cloning, it has been underdetermined in the fine mapping and cloning of insect resistance genes in  recent decades.”

 

                Underdetermined” is not the appropriate term

 

L376-377

 

“the potential function of these expressed sequences was likely  associated with the ability to maintain photosynthesis for tolerance to S. avenae”

 

Sitobion avenae is an aphid, it doesn’t reduce the photosynthesis of wheat…

 

L421-427

 

“dsRNA molecules should be sprayed multiple times during agricultural production. Meanwhile, the widespread and frequent application of sprayable dsRNA products targeting the crucial genes of pathogens or pests on the foliar of seedlings would significantly stimulate attackers to evolve RNAi resistance and increase the difficulty of pest control. Thus, the specific production of dsRNA derived from critical genes of different biotic stresses in transgenic plants may be a promising measure for future crop protection”

You explained 2 defaults of the dsRNA and you conclude (“thus”) that production of ds RNA may be promising ?? Producing dsRNA in the host plant can avoid to spray but the second default remains.. then … not promising at all.

 

 

L623-626

 

“…;Moreover, diverse attackers are more likely to colonize different or similar niches of the same plant to obtain nutrients, probably because their feeding or infestation could positively induce  biological and physiological alterations

Unclear, difficult to understand

 

L626-628

 

The crosstalk between phytohormone signaling pathways induced by multiple colonizers would fine-tune the plant defense re sponses and further aggravate the challenges of attackers

Unclear, difficult to understand

 

I did not get the 3 figures of your review...

 

Best regards,

 

Author Response

Dear Editor and Reviewers,

Thank you very much for your letter and the comments about our paper submitted to your journal. We would also like to thank the reviewers for their careful and constructive reviews. Soon after receiving your comments, the other authors and I carefully revised the manuscript. The changes made according to the comments from the reviewers are detailed below.

 

Reviewer #2: Please find some remarks and questions (en green) below:

Response 1: Thank you very much for carefully reviewing our manuscript and kindly giving us many suggestions and comments for improving the readability of this manuscript. In the revision, we carefully followed your corrections.

 

L27-28 :

“…production of bread wheat (Triticum aestivum L.) is of paramount importance to mankind, especially in arid and semiarid areas’’

The production of wheat is important in every area and not more in arid conditions… This is the first crop of the world.

 Response 2: Previously, we wanted to demonstrate the importance of wheat production worldwide. Although it occupies the largest cultivated area in the world, its grain yield is the third largest when compared with the yields of corn and rice. To improve the readability and logic of the Introduction, after careful consideration, we deleted some unrelated sentences and reworded some sentences to prepare a good Introduction.

 

 

L37-38:

 “…In addition, returning straw back into fields also potentially increases the rate of occurrence of plant diseases and insect pests in the subsequent season of crop growth…”

Personally, I think that this accurate information is not relevant in this part of the text, which present the general risk caused by biotic stress.

Response 3: We apologize for not using the accurate phase in that sentence. To improve the readability and logic of the Introduction, in the revision, we used the appropriate wording.

 

L42-43

            “…irrational application has considerably enhanced environmental issues and pest resurgence”

                Most of the application are not irrational in the whole world…

  Response 4: Previously, we wanted to demonstrate the phenomena of the widespread and frequent application of chemicals in farming, which is facilitated by pursuing high economic profits. Therefore, we hypothesize that the phase of “irrational application of chemicals” accurately describes our mind. To improve readability, in the revision, we used the appropriate wording.

L43-46

“… rapid increase in the consumption of high-quality foods and products derived from wheat grains, which has significantly promoted the development of wheat resistance and biofortification breeding programs.

There is no link between the increase in wheat consumption in some parts of the world and the development of resistant cultivars. Since the beginning of wheat breeding, breeders have tried to develop resistant and tolerant varieties.

 Response 5: We apologize for not using the accurate phase in that sentence. To improve the readability and logic of the Introduction, after careful consideration, we deleted these sentences to prepare a good Introduction. Please allow us to make the correction.

 

L62

“…Based on these advances, wheat breeders have given much attention to the development of hybrids…”

Be careful to define hybrids in wheat. By the past, ‘Hybrid” was synonymous with “crossing” but at the present time hybrid means Varieties in first generation sold to the farmer…

 

And as said before, breeders have not waited the molecular advances to give attention to the durable resistance to biotic attackers…

 Response 6: Thank you very much for the reword suggestion. We have replaced “hybrids” with “genotypes” in the revision. In addition, we rewrote that section and corrected the illogical points.

 

L66-69

“…Although some of these resistant wheat cultivars could adversely affect the performance of fungal diseases and herbivorous insects, there were still some obstacles that severely limited the large-scale practical  applications of resistant cultivars in agriculture.”

Are you talking about resistant cultivars obtained by new molecular techniques or by conventional breeding?

Response 7: Here, we wanted to mention that both approaches had been applied to the development of wheat cultivars in the past; however, there have still been some obstacles that have severely limited the large-scale practical applications of resistant cultivars in agriculture. Therefore, we summarize and discuss the recent advances and challenges related to breeding measures for improving common wheat cultivars with pest resistance and disease tolerance in the present review.

 

  Some problems in the first paragraph of section 2:

Response 8: We apologize that we did not stringently express the right meaning for the above statements in the original submission. This manuscript focused on characterizing the molecular advances in breeding for durable resistance against pests and diseases in wheat. After full consideration, we deleted the irrelevant section from the manuscript in the revision. Please allow us to delete this paragraph from the revision manuscript.

L76

“…by contrasting new… “

Appropriate term?

Response 9: As described previously, to improve the readability and logic of the Introduction, after careful consideration, we deleted these sentences to prepare a good Introduction. Please allow us to make the correction.

L84-85

                “….conventional plant breeding programs have always relied on phenotypic characteristics for the development of new cultivars or the improvement of existing cultivars…”

                Conventional breeding of wheat has always relied on phenotypic characteristics,  yes but also on quality of grains and flours analysis…

Response 10: It is true that the phenotypic selection in conventional wheat breeding programs always relies on grain quality and flour analysis.

L86-88

            “….breeders usually spend more time selecting wheat cultivars with the desired traits, while most of the preferred traits are controlled by quantitative trait loci (QTL)”

 

                I doubt that the “most preferred traits are the one of QTL instead of the one of the breeder.

 Response 11: Although some of the resistance traits characterized in wheat are monogenic and inherited as dominant traits, similar to chemical control, the practice of breeding for high levels of resistance often promotes the development of pest virulence. This strongly suggests the need to identify new and diverse resistance genes and genes that confer tolerance or more moderate levels of resistance in pest management, which attract more breeders to develop cultivars with tolerance or more moderate levels of resistance. However, these traits are always controlled by QTLs. Therefore, introducing these traits into elite cultivars would bring more difficulty to breeders.

 

L88-90

“…Therefore, against the new background of wheat breeding, conventional wheat breeding cannot fulfill the demands of sustainable agriculture.”

 

I strongly disagree with this statement! Please inform yourself on new varieties obtained all around the world by conventional breeder.

 Response 12: As described previously, to improve the readability and logic of the Introduction, after careful consideration, we deleted this sentence from section 2. In Figure 1, we compared the difference and the link between conventional wheat breeding programs and molecular marker-assisted breeding. This conclusion is supported by the evidence that conventional wheat breeding programs spend more time than molecular breeding programs. Additionally, many of the characterized resistant traits are always controlled by QTLs, which can be easily affected by diverse environmental factors. Introducing these traits into elite cultivars would bring more difficulty to breeders. Therefore, we want to express the challenge of conventional wheat breeding programs.

 

L92-93

“…could not only disrupt reproductive isolation”

 

This sentence is not clear

  Response 13: We are very sorry that we did not stringently express the right meaning for the above statements in the original submission. To improve the readability and logic of the Introduction, after careful consideration, we deleted this phase from section 2.

 

L99-100

                “…Thus, large-scale cultivation of GM wheat plants is considered to be a reliable and environmentally friendly approach to combat crop diseases and herbivorous insects in agricultural production.”

“Considered” by you, author? undoubtedly but what about the opinion of the rest of the world?

 

  Response 14: As described previously, molecular breeding of GM wheat plants could significantly reduce the process of developing resistant wheat when compared with conventional wheat breeding programs. The GM cultivars could easily transfer the desired traits and reduce undesired traits. According to this foundation and the conclusions from the references, we think large-scale cultivation of GM wheat plants is considered to be a reliable and environmentally friendly approach to combat crop diseases and herbivorous insects in agricultural production.

L125-128

“…Although there were some efforts focused on the development of transgenic wheat resistant to different cereal viral diseases until the early 2000s, the accumulating experimental evidence revealed that it is not a trivial task [23], probably because most viruses are transmitted by herbivorous insects”

 

             Do you mean that this work is trivial because of the aphids? Or should aphid transmission be considered an opportunity to stop the epidemic?

   Response 15: Yes, that is the important reason because the different serotypes of viruses are transmitted by different species of herbivorous insects; thus, we should pay more attention to developing cultivars with pest resistance. In that case, the population dynamics of pests could be significantly suppressed but could also alleviate the epidemics of viral diseases. In addition, most reported viral disease resistance genes or alleles do not confer tolerance to all serotypes of viruses. Thus, the development of transgenic wheat resistant to different cereal viral diseases is not a trivial task.

L151-L155

“…Meanwhile, a diverse set of foreign genes associated with the qualified traits of abiotic stress tolerance and yield and quality were integrated into elite wheat cultivars with ATMT and callus bombardment techniques, and the cultivars carrying these foreign genes exhibited high yield and quality and exhibited tolerance to different abiotic stresses.”

 

Could you give some example of elite cultivars obtained by these methods?

 Response 16: Yes, I can give examples of elite cultivars by strengthening their abiotic stress tolerance and yield and quality. For instance, expressing the barley HVA1 gene in wheat could improve its biomass productivity and wat use efficiency. (Sivamani E, Bahieldin A, Wraith J M, Al-Niemi T, Dyer W E, Ho T D H, Qu R. Improved biomass productivity and water use efficiency under water deficit conditions in transgenic wheat constitutively expressing the barley HVA1 gene. Plant Science, 2000, 155: 1-9.) However, the topic of this manuscript is focused on combining the different molecular-related approaches that have been implemented in the direction of breeding wheat varieties that are resistant or tolerant to different biotic stresses at the same time where opportunities exist and challenges thereon. Therefore, in the text, we did not illustrate examples of wheat cultivars carrying these foreign genes that exhibited high yield and quality and exhibited tolerance to different abiotic stresses.

L187

“DDuring”

 

Typing error

 Response 17: Thank you very much for pointing out the typing error. We have removed the extra “D”.

 

L216-L217

 

“…wheat is a natural hybrid of Triticum urartu (L. ), T. turgidum (L.) 216 and Aegilops tauschii (Coss. )”

Wheat is a natural hybrid of Triticum urartu (AA), a close parent of Aegilops speltoides (BB)) and Aegilops tauschii (DD). T. turgidum (AABB) is the product of the hybridization between the genomes AA and BB.

Response 18: Thank you very much for pointing out the error. We corrected that error at the end of section 4 in the revision of the manuscript.

 

L217-L219

 

“…it has the largest genome, probably because it has two copies of three groups of 42 chromosomes (2n = 6x = 42, genome formula AABBDD) obtained from each of its ancestors”

 

Each ancestor has not 42 chromosomes but only 2 x 7 chr. = 14 chr. Wheat has (3 x 14chr = 42chr.)

 Response 19: We apologize that we did not stringently express the right meaning for the above statements in the original submission. Here, we wanted to mention that common wheat plants have an extremely large genome size (16,000 Mbp). After full consideration, we deleted the irrelevant information from the manuscript in the revision. Please allow us to delete this sentence from the revision manuscript.

 

L231-233

 

“….QRp.slu.4BL, which conferred antibiosis resistance against the bird cherry-oat aphid (Rhopalosiphum padi L. ), while QRp.slu.5AL and QRp.slu.5BL conferred tolerance to R. padi

? then all the 3 conferred antibiosis resistance to R. padi … why the author use “while”?

 Response 20: These three QTLs conferred distinct resistance mechanisms. Therefore, we use the “while” to express the difference. We are very sorry that we did not stringently express the right meaning for the above statements in the original submission. In the revision, we rewrote that section and corrected the illogical points in the conclusion.

 

L280-281

 

“…Taken together, these molecular markers involved in MAS would enable a reliable and feasible approach to detect resistant germplasms and track resistance genes in wheat breeding programs.

 

Finding gene mutations does not mean finding resistance. Most of the mutations do not have the expected effect on the phenotype in the field. The most “reliable” means to detect resistance is to observe the plant in their environment when they face (a)biotic stress…

  Response 21: Yes, it is true. We are very sorry that we did not stringently express the right meaning for the above statements in the original submission. After full consideration, we deleted the irrelevant information from the manuscript in the revision. Please allow us to delete this sentence from the revision manuscript.

 

L298-302

“Moreover, common (hexaploid) wheat is a natural hybrid of Triticum urartu (L. ), 298 T. turgidum (L.) and Aegilops tauschii (Coss. ), and it has an extremely large genome size 299 (16,000 Mbp), probably because it has two copies of three groups of 42 chromosomes (2n 300 = 6x = 42, genome formula AABBDD) obtained from each of its ancestors and encodes an 301 estimated 164,000 to 334,000 genes [38].”

 

This sentence is the exact repetition of the line L216-L219…

 Response 22: Thank you very much for pointing out the repetitive error. We have removed this part in the revision.

L369-371

 

                “In comparison with disease resistance gene cloning, it has been underdetermined in the fine mapping and cloning of insect resistance genes in  recent decades.”

 

“Underdetermined” is not the appropriate term

 Response 23: Thank you very much for the reword suggestion. We have rewritten that sentence and corrected the illogical points.

L376-377

 

“the potential function of these expressed sequences was likely associated with the ability to maintain photosynthesis for tolerance to S. avenae”

 

Sitobion avenae is an aphid, it doesn’t reduce the photosynthesis of wheat…

 Response 24: Aphids do not reduce the photosynthesis of wheat; however, they can absorb photoassimilates from plant tissues. Therefore, the tolerant wheat plants increase the photosynthesis of wheat to convert additional photoassimilates to compensate for the damage caused by aphid feeding and strengthen the cell wall for tolerance to aphid infestation. We are very sorry that we did not stringently express the right meaning for the above statements in the original submission. To improve its readability and logic, we have rewritten that sentence and corrected the illogical points.

L421-427

 

“dsRNA molecules should be sprayed multiple times during agricultural production. Meanwhile, the widespread and frequent application of sprayable dsRNA products targeting the crucial genes of pathogens or pests on the leaves of seedlings would significantly stimulate attackers to evolve RNAi resistance and increase the difficulty of pest control. Thus, the specific production of dsRNA derived from critical genes of different biotic stresses in transgenic plants may be a promising measure for future crop protection.”

You explained 2 defaults of the dsRNA and you conclude (“thus”) that production of ds RNA may be promising?? Producing dsRNA in the host plant can avoid spraying, but the second default remains.. then … not promising at all.

Response 25:  We’re very sorry for we didn’t stringently express the right meaning for the above statements in the original submission. The second default should be that producing dsRNA in the host plant can avoid the attackers to evolve RNAi resistance. Because it could more accurately to control the pests at the correct timepoint, and reduce the drawbacks of spraying, such as reduce the chance of diffusion the dsRNA in the environment, and it also reduce the amount of dsRNA from the frequency.

 

 

 

L623-626

 

“…Moreover, diverse attackers are more likely to colonize different or similar niches of the same plant to obtain nutrients, probably because their feeding or infestation could positively induce biological and physiological alterations

Unclear, difficult to understand

  Response 26: In agroecosystems, it is common for diverse attackers to colonize different areas; for instance, the phloem feeders S. avenae and F. graminearum are two economically important pests residing on wheat spikes. Our previous manuscript established the working hypotheses of ‘synergistic’ effects of the coexistence of aphids and phytopathogens on wheat spikes in agroecosystems from the aspects of environmental nutrients, auxin production, hormone signals, and their potential roles.

L626-628

 

The crosstalk between phytohormone signaling pathways induced by multiple colonizers fine-tunes plant defense responses and further aggravates the challenges of attackers

Unclear, difficult to understand

 Response 27: Once plant invaders successfully colonize and begin feeding, plants trigger induced defenses to counter the effects of their attack. The induced defenses are always mediated through the release of phytohormones that specifically activate hormone-dependent response pathways against various attackers. The signaling molecules jasmonic acid (JA) and salicylic acid (SA) are recognized as major defense hormones. These phytohormone signals trigger distinct defense responses in the species of attackers. JA-mediated defense responses are always associated with resistance to necrotrophic pathogens, whereas the direct and indirect defense responses mediated by SA are the most important plant defenses against biotrophic and hemibiotrophic pathogenic fungi. Accumulating experimental evidence has demonstrated that SA and JA signaling cascades in plant immune responses always act coordinately; in most cases, antagonistic interactions between the two signaling pathways have been detected in diverse fungi-plant systems. Our previous manuscript established the working hypotheses of ‘synergistic’ effects of the coexistence of aphids and phytopathogens on wheat spikes in agroecosystems, from the aspects of hormone signals to fine-tune the plant defenses to benefit them.

I did not get the 3 figures of your review...

 Response 28: We apologize for not uploading the three figures in the original submission. To improve the readability and logic of the figures, in this manuscript, we modified the text and the arrows in the figures and added more content to the descriptions in the figure legends following your suggestion and comments.

Author Response File: Author Response.docx

Reviewer 3 Report

The paper contains comprehensive review on the molecular work done so far on wheat breeding and opportunities to explore are indicated as well as the challenges that are likely to be faced in implementing some of the modern technologies to improve breeding of biotic and abiotic stress tolerant durum wheat varieties

-Title need to correct pest and disease to read “Pests and Diseases”

Line 187- correct DDuring to read During

-There seems to be a repeat of same statements from Line 214-Line 221 and the statements in line 298-Line 305 revisit

-Line 370 I feel the phrase insect resistance genes should read “insect resistant genes”

-Line 429 there is reference made to Figure 2 and am wondering if there is Figure 1 somewhere and l didn’t the figure being referred to

-Line 607 reference is made to Figure 3 can the figure be included unless the figures were in a separate document that l did not access

 

More comments:

1) What is the main question addressed by the research?

The research is a review that has tried to put together the different molecular related approaches that have been implemented in the direction of breeding wheat varieties that are tolerant to biotic and abiotic stresses at the same time highlighting where opportunities exist and challenges thereon

2) Do you consider the topic original or relevant in the field? Does it address a specific gap in the field? 

The topic is relevant in the field and the gap it is addressing is that of integrating the different interventions to come up with durable resistance that will see very good wheat hybrids being produced to address food security issues. The topic can be modified to read

“Molecular advances in breeding for durable resistance against pests and diseases in wheat: opportunities and challenges”

3) What does it add to the subject area compared with other published material

The paper is complementing the work already done by other scientists by highlighting and emphasizing the need for intensification of efforts to come up with novel wheat varieties.

4) What specific improvements should the authors consider regarding the methodology? What further controls should be considered?

a) Revisit the introduction giving more insight into trait improvement that has taken place in wheat varieties because of molecular related interventions

b) Breeding aspects can be given a clear flow chart of integration of these genomics/genes into breeding pipelines 

5) Are the conclusions consistent with evidence and arguments presented and do they address the main question posed?

The conclusions don’t seem to be consistent with evidence in the methodology the issue of challenges with use of transgenics was discussed but in the prospects the authors are silent concerning the wayforward as far as this technology is concerned.

6) Are the references appropriate

The refences are appropriate

7) Comments on Tables and figures

Reference is made to figures in the methodology but l did not come across any graph maybe l might have missed these ones

Author Response

Dear Editor and Reviewers,

Thank you very much for your letter and the comments about our paper submitted to your journal. We would also like to thank the reviewers for their careful and constructive reviews. Soon after receiving your comments, the other authors and I carefully revised the manuscript. The changes made according to the comments from the reviewers are detailed below.

 

Reviewer #3:

The paper contains a comprehensive review of the molecular work done thus far on wheat breeding, and opportunities to explore are indicated as well as the challenges that are likely to be faced in implementing some of the modern technologies to improve the breeding of biotic and abiotic stress-tolerant durum wheat varieties.

-Title need to correct pest and disease to read “Pests and Diseases”

Response 1: We’re very sorry for we didn’t prepare a suitable title in the original submission. Thank you very much for the title suggestion. We prepared a new title for the revision. We guess this title is more suitable for the content of this manuscript.

Line 187- correct DDuring to read During

Response 2: Thank you very much for pointing out the typing error. We have removed the extra “D”.

-There seems to be a repeat of same statements from Line 214-Line 221 and the statements in line 298-Line 305 revisit

-Line 370 I feel the phrase insect resistance genes should read “insect resistance genes”

Response 3: Thank you very much for the suggestion. We have replaced “insect resistance genes” with “insect resistance genes” in the revision.

 

-Line 429 there is reference made to Figure 2 and am wondering if there is Figure 1 somewhere and l didn’t the figure being referred to

Response 4: We apologize for not uploading the three figures in the original submission. Figure 1 illustrates the regular flow charts of the integration of desirable traits into wheat cultivars in wheat breeding programs, and Figure 2 illustrates prevalent wheat germplasm improvement strategies and the principal stages in genetic engineering in wheat breeding.

-Line 607 reference is made to Figure 3 can the figure be included unless the figures were in a separate document that l did not access

Response 5: We apologize for not uploading the three figures in the original submission. To improve the readability and logic of the figures, in this manuscript, we modified the text and the arrows in the figures and added more content to the descriptions in the figure legends.

 

More comments:

1) What is the main question addressed by the research?

The research is a review that has tried to put together the different molecular-related approaches that have been implemented in the direction of breeding wheat varieties that are tolerant to biotic and abiotic stresses at the same time highlighting where opportunities exist and challenges thereon.

2) Do you consider the topic original or relevant in the field? Does it address a specific gap in the field?

The topic is relevant in the field and the gap it is addressing is that of integrating the different interventions to come up with durable resistance that will see very good wheat hybrids being produced to address food security issues. The topic can be modified to read

“Molecular advances in breeding for durable resistance against pests and diseases in wheat: opportunities and challenges”

Response 6: We’re very sorry for we didn’t prepare a suitable title in the original submission. Thank you very much for the title suggestion. We prepared a new title for the revision. We guess this title is more suitable for the content of this manuscript.

3) What does it add to the subject area compared with other published material

The paper complements the work already done by other scientists by highlighting and emphasizing the need for intensification of efforts to develop novel wheat varieties.

4) What specific improvements should the authors consider regarding the methodology? What further controls should be considered?

  1. a) Revisiting the introduction, giving more insight into trait improvement that has taken place in wheat varieties because of molecular-related interventions

Response 7: To improve the readability and logicality of the introduction section in this manuscript, we have rewritten the introduction strictly following your suggestion and comments. Please allow us to make those revisions.

 

  1. b) Breeding aspects can be given a clear flow chart of the integration of these genomics/genes into breeding pipelines.

Response 8: Yes, the present review has focused on combining the different molecular-related approaches that have been implemented in the direction of breeding wheat varieties that are resistant or tolerant to different biotic stresses at the same time where opportunities exist and challenges exist. Thus, we prepared three figures to illustrate the regular flow charts of the integration of desirable traits into wheat cultivars in wheat breeding programs, prevalent wheat germplasm improvement strategies and the principal stages in genetic engineering in wheat breeding, and a model summarizing the approaches for future development of wheat cultivars with durable improvements. We hypothesize that these three figures could more accurately describe the recent advances and drawbacks related to the measures associated with molecular breeding in common wheat cultivar improvement for the readers.

5) Are the conclusions consistent with evidence and arguments presented and do they address the main question posed?

The conclusions don’t seem to be consistent with evidence in the methodology the issue of challenges with use of transgenics was discussed but in the prospects the authors are silent concerning the way forward as far as this technology is concerned.

Response 9: It’s true that we didn’t concerning more on the challenges of the transgenic wheat breeding programs. However, we had discussed this topic on the section 2, about the health, ecological, and environmental issues. The accumulating evidence demonstrated that identify new and diverse resistance genes and genes that confer tolerance or more moderate levels of resistance in pest management, which attract more breeders to develop the cultivars with tolerance or more moderate levels of resistance. In comparisons with transferring the foreign genes from different species, transferring the inherent genes from resistant wheat cultivars, or silencing or editing the susceptible genes in wheat plant would provide the alternative ways to breeding novel cultivars. Thus, in the prospects section, we focused more on this direction. We guess it could represent the tendency of wheat breeding programs.

6) Are the references appropriate

The refences are appropriate

7) Comments on Tables and figures

Reference is made to figures in the methodology but l did not come across any graph maybe l might have missed these ones

Response 10: We apologize for not uploading the three figures in the original submission. To improve the readability and logic of the figures, in this manuscript, we modified the text and the arrows in the figures and added more content to the descriptions in the figure legends.

 

Author Response File: Author Response.docx

Round 2

Reviewer 1 Report

Luo et al. revised review manuscript entitled “Molecular Advances in Breeding for Durable Resistance against Pests and Diseases in Wheat: Opportunities and Challenges.”  Authors considered the reviewers suggestions and improved the review manuscript in the revised version.

Author Response

Dear Reviewer,

Thank you very much for your letter and the comments about our paper submitted to your journal. We would also like to thank you for your careful and constructive reviews. Soon after receiving your comments, the other authors and I carefully revised the manuscript. The changes made according to the comments from the academic editor are detailed below.

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