the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
| 28 Jun 2024
A new approach to continuous monitoring of carbon use efficiency and biosynthesis in soil microbes from measurement of CO2 and O2
Abstract. Soils comprise the largest terrestrial carbon pool. Therefore, understanding processes that control soil carbon stabilization and release is vital to improving our understanding of the global carbon cycle. Heterotrophic respiration is the main pathway by which soil organic carbon is returned to the atmosphere, however not all carbon utilized by heterotrophs shares this fate, as some portion is retained in the soil as biomass and biosynthesized extracellular compounds. The fraction of carbon consumed by microbes that is used for biomass growth (the carbon use efficiency or CUE) is an important variable controlling soil carbon stocks but is difficult to measure. Here we show that CUE can be continuously monitored in laboratory glucose-amended soil incubations by measuring CO2 and O2 gas concentrations, allowing instantaneous estimates of microbial biomass growth. We derive a theoretical relationship between the respiratory quotient (RQ), the ratio of carbon dioxide produced to oxygen consumed during respiration, and CUE that recognizes the influence of both substrate and biosynthesized product oxidation states on RQ. Assuming the biosynthesized product has the stoichiometry of an average microbe, and that the substrate is primarily the glucose used for amendment, we measure RQ and use our theoretical relationship to calculate CUE, and from that, biomass production. Extractions of microbial biomass carbon at the end of the experiments reveal minimal net increases in standing biomass across all amended treatments, suggesting that much of this newly produced biomass is likely converted to necromass as substrate availability declines and this results in a net storage of new soil organic matter. Carbon budgets compiled from measurements of relevant pools account for the amended carbon and suggest that with larger carbon amendments, increases in C : N ratios lead to increases in the relative portion of the amendment acutely lost from the soil. These findings demonstrate that soil RQ values may be used to monitor changes in CUE and that studies which monitor soil RQ values should consider CUE as a key factor when changes in RQ are observed, for instance, with changing environmental conditions or changes in production of plant derived compounds. This new approach may be leveraged to provide information on the storage of soil organic matter. These findings demonstrate how measurements of soil RQ may be leveraged to understand soil carbon transformations, specifically the fate of fresh carbon inputs.
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Status: closed
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RC1: 'Comment on egusphere-2024-1757', Anonymous Referee #1, 01 Jul 2024
The reviewed manuscript presents a new approach to the measurement of CUE. Given the importance of CUE for carbon preservation in soils, such a suggestion for a fast and automated approach is intriguing. The calculations connecting RQ to CUE the authors present are straightforward. However, as noted in line 83 there are other controls on RQ including "calcite dissolution/precipitation which can cause a transient decoupling of these two gases, and oxidation of reduced metal species". In addition, anaerobic respiration will cause an increase in the RQ, and oxygen-poor microsites can be found in well-aerated soils. This fact is not noted in the manuscript. This is a major caveat of the suggested approach which requires extensive discussion. It may pull the rug under the entire section 4.3.
I am also missing a comparison of this experiment with experiments measuring CUE using other methods. Is an increase in CUE with added glucose common? Or maybe the peak in RQ at the same time as the peak in respiration indicates that high O2 consumption rates led to local anaerobic conditions?
Minor comments:
Line 36: use of "around" followed by a 4-digits-accuracy number is awkward.
Line 82: The sentence there needs re-phrasing. Something like:
other studies observed systematic deviations that were linked to nonmetabolic processes that can affect the soil CO2 and O2 fluxes, and thus noted the ratio of these fluxes as the Apparent Respiratory Quotient (ARQ). Such nonmetabolic processes include…
Also, add here the role of anaerobic respiration.
Line 94: Better to just write every 2 hours. High temporal resolution can mean anything from seconds to days.
Line 265: " It is important to note that once RQ values drop below a value 1.0, the modeled RQ—CUE relationship for glucose as the sole substrate no longer applies."
This is confusing: So until RQ=1.0 the model works and then suddenly it is not applicable? Most likely the same processes causing a RQ<1.0 that the model cannot account for, are already in play before that. So is the model relevant at all? There is a need to expand on this.
Line 299-301: Please rephrase. I could not understand this long sentence.
In general, the paper is well-written and clear.
Citation: https://doi.org/10.5194/egusphere-2024-1757-RC1 -
AC1: 'Reply on RC1', Kyle Smart, 12 Jul 2024
We thank the referee for their suggestions and their perspective. We would like to first address the concern about the presence of anaerobic respiration and its implications on our findings.
We agree that if meaningful amounts of anaerobic respiration occurred, that could drive RQ values greater than 1.0. However, there are several factors that lead us to conclude anaerobic respiration is not the primary factor driving the elevated RQ values we measured, and these are listed as follows:
1) Soil headspace gas measurements were measured throughout the incubation, and even at peak respiration oxygen concentrations remained above 19% (these measurements can be found in the zenodo data repository).
2) The soil existed as only a thin layer within our bottles (no more than 1cm thick) and was comprised of particles <2mm in size. Given the strong inhibitory nature of oxygen on anaerobic respiration, we believe this would support abundant aerobic respiration and limit any anaerobic respiration. Further, the effect of sampling headspace and returning CO2-Free Air should effectively drive atmospheric circulation in the bottle.
3) Anaerobic respiration tends to occur more slowly than aerobic respiration, which further limits its possible impacts on our measurements.
4) The close agreement in the carbon budget presented in figure 5 suggests that aerobic respiration was driving the elevated RQ values we observed. Otherwise, during peak respiration ~50 % of the CO2 produced would have been produced through anaerobic respiration, meaning both anaerobic and aerobic respiration would be matched and this is not likely possible given incubation conditions.
5) To account for RQ values around 1.5 like we observe, anaerobic respiration rates would need to be significant during peak respiration, when oxygen concentrations were still greater than 19%.
We acknowledge that we cannot entirely rule out the presence of any anaerobic respiration from having occurred; however, the data and the controlled incubation conditions imply that it was not significant in these incubations.
To address the second larger concern, the soil pre-treatment as described in line 136, was intentional to place the soil microbes in a stressed state with lower amounts of standing biomass so that when re-wet and amended with glucose, this would stimulate the production of new biomass in effort to measure its effects on RQ. This stimulation of biomass growth would require that bulk soil CUE would be greater than 0. CUE at peak respiration of around 0.4 is generally in line with other works which commonly range from 0.4-0.6. Many of these works, amended glucose to a relatively undisturbed soil in comparison to our soil pre-treatment which makes comparing them precarious. Though it is documented that with changes in environmental conditions, such as available substrate and soil moisture, changes in CUE do occur.
Changes in the text to address comments:
Line 36 - removed the word “around”.
Line 82 - sentence has been reworked to address ARQ first, then discuss the other non-metabolic processes, and added sentences addressing anaerobic respiration.
Line 94 - “High temporal resolution” removed.
Lines 168 - 171 - Added statement regarding experimental design with intent to prevent and/or minimize anaerobic respiration from impacting our incubation results.
Lines 273-275 - Follow up sentence, stating why this relationship can no longer be applied when RQ drops below 1.0, and what is likely occurring when this happens.
Lines 287-289- Added sentence discussing expectations for temporally dynamic CUE.
Lines 291-293- Added sentence discussing alternative explanations for RQ values and CUE as RQ values transition from >1.0 to <1.0.
Lines 311-313 - have been re-written more concisely for clarity.
Lines 319-320 - extended the sentence to mention that close agreement in Figure 5 suggests that anaerobic respiration was not likely an important process in these incubations.
Changes have been made to the document and will be updated with the next submission.
Citation: https://doi.org/10.5194/egusphere-2024-1757-AC1
-
AC1: 'Reply on RC1', Kyle Smart, 12 Jul 2024
-
RC2: 'Comment on egusphere-2024-1757', Xianjin He, 08 Jul 2024
General Comments:
The manuscript "A new approach to continuous monitoring of carbon use efficiency and biosynthesis in soil microbes from measurement of CO2 and O2" presents a novel methodology for monitoring microbial CUE through measuring the Respiratory Quotient (RQ). The study is interesting, especially the authors providing a theoretical link between RQ and CUE. The experiments and data analysis are robust. The findings have some useful implications for understanding soil carbon dynamics and microbial metabolism.
I hope the authors can supplement the revised manuscript in the following two aspects:
First, I hope the authors can clarify the specific application prospects of this new method. Currently, there are already various methods for testing CUE, and it is difficult to unify them. This study indeed provides a novel approach. However, I did not read from the text what scientific problems this new method can be used to solve. Additionally, the authors emphasize that this method can get the temporal dynamics of CUE, as shown in figure 4b. But I think this is also a confusing point of this method: if CUE changes with incubation time, then which time point's value can be used to represent the microbial CUE of this soil sample? Or should it be the average value over a certain period? The answer to this question relates to what scientific problems this method can be used to solve. I hope the authors can supplement their thoughts or suggestions in this regard.
Second, this method is based on a mass conservation formula of C, H, O, and N elements to derive the relationship between RQ and CUE. This formula is a simplification of the real ecosystem. I hope the authors can discuss the limitations or deficiencies of this method to help future researchers using this method understand its limitations.
Specific Comments:
1 Introduction
The introduction is well-written and reads smoothly. However, I noticed multiple formatting errors in the references, e.g., in lines 49, 54, 56, 59, 68, etc.. Please carefully check the formatting of the authors' names in the references.
2 Connecting Carbon Use Efficiency and Respiratory Quotient
I suggest moving the content from the appendix to the main text. The information in the appendix is essential for understanding how the relationship between CUE and RQ is derived. Additionally, in the appendix, the derivation from step 1 and 2 to step 3 is not straightforward. Could you provide a more detailed derivation process?
3 Materials and Methods
Line 135: I am curious why glucose was added as a fine solid powder instead of being dissolved in water and then added to the soil. Was there a specific reason for this choice?
4 Results and Discussion
In lines 222, 223, 232, and 233, the figure numbers were missed.
Line 265, from the results in figure 4b, it appears that only a small portion of the observations from the 100 mg treatment can be used to calculate CUE. Does this imply that larger doses of glucose should be considered when using this method to measure CUE? I suggest discussing this point.
Figure 4: Why are the results of only one replicate presented instead of the average values of all replicates? If only one replicate result is shown, how did you select which replicate to present from among the several replicates? Please explain.
Citation: https://doi.org/10.5194/egusphere-2024-1757-RC2 -
AC2: 'Reply on RC2', Kyle Smart, 21 Jul 2024
We thank the referee for their thoughtful comments and suggestions.
We believe that the applications of this method and relevant timescale of application are intimately linked questions. The exciting aspect of this approach is the relatively high resolution of these CUE estimates, which could be used to bolster our understanding of fast responses within the microbial community to environmental changes without the time intensive laboratory work associated with other methods for measurement of CUE. The other exciting aspect of this approach is that the metabolism in question can be tailored to answer different types of questions by using the same type of reaction-derivation, this can be applied to many other specific substrates. The second key aspect of this work is that we validate a theoretical link between CUE and measured RQ values. This knowledge will help provide context for field studies of RQ as well, now with the knowledge that transient shifts may be a result of microbial processes separate from changing substrates.
We believe this method will enable the observation of CUE at resolutions that would be considered logistically challenging with other methods, while the incubations we carry out are done with relative ease after the construction of the apparatus has taken place. We believe the types of scientific questions with which this method would provide strong advantage is the centered on this temporal aspect, for example studying the effects of dynamic environmental conditions on microbial metabolisms and carbon use efficiencies. Further, these incubations could be carried out for extended periods of time (several weeks to months) and provide relatively high resolution on not only CUE when substrate is well constrained, but for example study other processes which are predicted/known to impact measurements of RQ. With minor modifications, this setup could be used in application to measure soil microbial respiratory behavior as oxygen is used up and conditions begin shifting to anaerobic, in effort to better understand the sensitivity and contribution of anaerobic respiration to bulk soil respiration.
Our thoughts are a single CUE that would best represent this period of glucose driven respiration would be a respiration weighted average of CUE over the period when RQ is ≥ 1. We believe this method is the most appropriate representation of CUE of the soil microbes specific to the period of time when glucose is the primary substrate.
We acknowledge that only considering C, H, O, and N is indeed a simplification of the real ecosystem and we hope that this work serves as a stepping stone and proof of concept. However, we are assured from Figure 5, that this fairly succinct simplification does effectively capture the process of biosynthesis in a manner which is approachable and compares very favorably with a carbon budget. However, further work could be done to provide an even more descriptive reaction to derive a more complete stoichiometric representation of soil microbes. However, it is also important to remember that other work has shown that microbial stoichiometry can vary with environmental conditions, so a more precise stoichiometry may be more applicable in certain use cases, but not all. Additional elements that could be included in this are phosphorus or sulfur, however their stoichiometric contributions are quite small in comparison to C, H, O, and N.
Comments Addressed:
Multiple lines : fixed in text citations.
Line 110, 298: reformatted Equations.
Lines 113 - 139: Moved appendix reaction derivation and added more derivation steps for readers to follow.
Lines 167 - 169 : Added sentence justifying our choice to add glucose as solid fine powder.
Lines 257, 258, 267-269: Added in figure numbers.
Line 329 : Added in reason for showing one replicate and method of choosing replicate, and why.
Lines 428 - 439: Added statements about how we think a prescribed CUE for the soils should be calculated and discussed the limitation of only using CHON elements to represent the system, as well as which amendment sizes are recommended for further experiments and future applications of this research.
Changes have been made to the working document and will be included in the next upload.
Citation: https://doi.org/10.5194/egusphere-2024-1757-AC2
-
AC2: 'Reply on RC2', Kyle Smart, 21 Jul 2024
Status: closed
-
RC1: 'Comment on egusphere-2024-1757', Anonymous Referee #1, 01 Jul 2024
The reviewed manuscript presents a new approach to the measurement of CUE. Given the importance of CUE for carbon preservation in soils, such a suggestion for a fast and automated approach is intriguing. The calculations connecting RQ to CUE the authors present are straightforward. However, as noted in line 83 there are other controls on RQ including "calcite dissolution/precipitation which can cause a transient decoupling of these two gases, and oxidation of reduced metal species". In addition, anaerobic respiration will cause an increase in the RQ, and oxygen-poor microsites can be found in well-aerated soils. This fact is not noted in the manuscript. This is a major caveat of the suggested approach which requires extensive discussion. It may pull the rug under the entire section 4.3.
I am also missing a comparison of this experiment with experiments measuring CUE using other methods. Is an increase in CUE with added glucose common? Or maybe the peak in RQ at the same time as the peak in respiration indicates that high O2 consumption rates led to local anaerobic conditions?
Minor comments:
Line 36: use of "around" followed by a 4-digits-accuracy number is awkward.
Line 82: The sentence there needs re-phrasing. Something like:
other studies observed systematic deviations that were linked to nonmetabolic processes that can affect the soil CO2 and O2 fluxes, and thus noted the ratio of these fluxes as the Apparent Respiratory Quotient (ARQ). Such nonmetabolic processes include…
Also, add here the role of anaerobic respiration.
Line 94: Better to just write every 2 hours. High temporal resolution can mean anything from seconds to days.
Line 265: " It is important to note that once RQ values drop below a value 1.0, the modeled RQ—CUE relationship for glucose as the sole substrate no longer applies."
This is confusing: So until RQ=1.0 the model works and then suddenly it is not applicable? Most likely the same processes causing a RQ<1.0 that the model cannot account for, are already in play before that. So is the model relevant at all? There is a need to expand on this.
Line 299-301: Please rephrase. I could not understand this long sentence.
In general, the paper is well-written and clear.
Citation: https://doi.org/10.5194/egusphere-2024-1757-RC1 -
AC1: 'Reply on RC1', Kyle Smart, 12 Jul 2024
We thank the referee for their suggestions and their perspective. We would like to first address the concern about the presence of anaerobic respiration and its implications on our findings.
We agree that if meaningful amounts of anaerobic respiration occurred, that could drive RQ values greater than 1.0. However, there are several factors that lead us to conclude anaerobic respiration is not the primary factor driving the elevated RQ values we measured, and these are listed as follows:
1) Soil headspace gas measurements were measured throughout the incubation, and even at peak respiration oxygen concentrations remained above 19% (these measurements can be found in the zenodo data repository).
2) The soil existed as only a thin layer within our bottles (no more than 1cm thick) and was comprised of particles <2mm in size. Given the strong inhibitory nature of oxygen on anaerobic respiration, we believe this would support abundant aerobic respiration and limit any anaerobic respiration. Further, the effect of sampling headspace and returning CO2-Free Air should effectively drive atmospheric circulation in the bottle.
3) Anaerobic respiration tends to occur more slowly than aerobic respiration, which further limits its possible impacts on our measurements.
4) The close agreement in the carbon budget presented in figure 5 suggests that aerobic respiration was driving the elevated RQ values we observed. Otherwise, during peak respiration ~50 % of the CO2 produced would have been produced through anaerobic respiration, meaning both anaerobic and aerobic respiration would be matched and this is not likely possible given incubation conditions.
5) To account for RQ values around 1.5 like we observe, anaerobic respiration rates would need to be significant during peak respiration, when oxygen concentrations were still greater than 19%.
We acknowledge that we cannot entirely rule out the presence of any anaerobic respiration from having occurred; however, the data and the controlled incubation conditions imply that it was not significant in these incubations.
To address the second larger concern, the soil pre-treatment as described in line 136, was intentional to place the soil microbes in a stressed state with lower amounts of standing biomass so that when re-wet and amended with glucose, this would stimulate the production of new biomass in effort to measure its effects on RQ. This stimulation of biomass growth would require that bulk soil CUE would be greater than 0. CUE at peak respiration of around 0.4 is generally in line with other works which commonly range from 0.4-0.6. Many of these works, amended glucose to a relatively undisturbed soil in comparison to our soil pre-treatment which makes comparing them precarious. Though it is documented that with changes in environmental conditions, such as available substrate and soil moisture, changes in CUE do occur.
Changes in the text to address comments:
Line 36 - removed the word “around”.
Line 82 - sentence has been reworked to address ARQ first, then discuss the other non-metabolic processes, and added sentences addressing anaerobic respiration.
Line 94 - “High temporal resolution” removed.
Lines 168 - 171 - Added statement regarding experimental design with intent to prevent and/or minimize anaerobic respiration from impacting our incubation results.
Lines 273-275 - Follow up sentence, stating why this relationship can no longer be applied when RQ drops below 1.0, and what is likely occurring when this happens.
Lines 287-289- Added sentence discussing expectations for temporally dynamic CUE.
Lines 291-293- Added sentence discussing alternative explanations for RQ values and CUE as RQ values transition from >1.0 to <1.0.
Lines 311-313 - have been re-written more concisely for clarity.
Lines 319-320 - extended the sentence to mention that close agreement in Figure 5 suggests that anaerobic respiration was not likely an important process in these incubations.
Changes have been made to the document and will be updated with the next submission.
Citation: https://doi.org/10.5194/egusphere-2024-1757-AC1
-
AC1: 'Reply on RC1', Kyle Smart, 12 Jul 2024
-
RC2: 'Comment on egusphere-2024-1757', Xianjin He, 08 Jul 2024
General Comments:
The manuscript "A new approach to continuous monitoring of carbon use efficiency and biosynthesis in soil microbes from measurement of CO2 and O2" presents a novel methodology for monitoring microbial CUE through measuring the Respiratory Quotient (RQ). The study is interesting, especially the authors providing a theoretical link between RQ and CUE. The experiments and data analysis are robust. The findings have some useful implications for understanding soil carbon dynamics and microbial metabolism.
I hope the authors can supplement the revised manuscript in the following two aspects:
First, I hope the authors can clarify the specific application prospects of this new method. Currently, there are already various methods for testing CUE, and it is difficult to unify them. This study indeed provides a novel approach. However, I did not read from the text what scientific problems this new method can be used to solve. Additionally, the authors emphasize that this method can get the temporal dynamics of CUE, as shown in figure 4b. But I think this is also a confusing point of this method: if CUE changes with incubation time, then which time point's value can be used to represent the microbial CUE of this soil sample? Or should it be the average value over a certain period? The answer to this question relates to what scientific problems this method can be used to solve. I hope the authors can supplement their thoughts or suggestions in this regard.
Second, this method is based on a mass conservation formula of C, H, O, and N elements to derive the relationship between RQ and CUE. This formula is a simplification of the real ecosystem. I hope the authors can discuss the limitations or deficiencies of this method to help future researchers using this method understand its limitations.
Specific Comments:
1 Introduction
The introduction is well-written and reads smoothly. However, I noticed multiple formatting errors in the references, e.g., in lines 49, 54, 56, 59, 68, etc.. Please carefully check the formatting of the authors' names in the references.
2 Connecting Carbon Use Efficiency and Respiratory Quotient
I suggest moving the content from the appendix to the main text. The information in the appendix is essential for understanding how the relationship between CUE and RQ is derived. Additionally, in the appendix, the derivation from step 1 and 2 to step 3 is not straightforward. Could you provide a more detailed derivation process?
3 Materials and Methods
Line 135: I am curious why glucose was added as a fine solid powder instead of being dissolved in water and then added to the soil. Was there a specific reason for this choice?
4 Results and Discussion
In lines 222, 223, 232, and 233, the figure numbers were missed.
Line 265, from the results in figure 4b, it appears that only a small portion of the observations from the 100 mg treatment can be used to calculate CUE. Does this imply that larger doses of glucose should be considered when using this method to measure CUE? I suggest discussing this point.
Figure 4: Why are the results of only one replicate presented instead of the average values of all replicates? If only one replicate result is shown, how did you select which replicate to present from among the several replicates? Please explain.
Citation: https://doi.org/10.5194/egusphere-2024-1757-RC2 -
AC2: 'Reply on RC2', Kyle Smart, 21 Jul 2024
We thank the referee for their thoughtful comments and suggestions.
We believe that the applications of this method and relevant timescale of application are intimately linked questions. The exciting aspect of this approach is the relatively high resolution of these CUE estimates, which could be used to bolster our understanding of fast responses within the microbial community to environmental changes without the time intensive laboratory work associated with other methods for measurement of CUE. The other exciting aspect of this approach is that the metabolism in question can be tailored to answer different types of questions by using the same type of reaction-derivation, this can be applied to many other specific substrates. The second key aspect of this work is that we validate a theoretical link between CUE and measured RQ values. This knowledge will help provide context for field studies of RQ as well, now with the knowledge that transient shifts may be a result of microbial processes separate from changing substrates.
We believe this method will enable the observation of CUE at resolutions that would be considered logistically challenging with other methods, while the incubations we carry out are done with relative ease after the construction of the apparatus has taken place. We believe the types of scientific questions with which this method would provide strong advantage is the centered on this temporal aspect, for example studying the effects of dynamic environmental conditions on microbial metabolisms and carbon use efficiencies. Further, these incubations could be carried out for extended periods of time (several weeks to months) and provide relatively high resolution on not only CUE when substrate is well constrained, but for example study other processes which are predicted/known to impact measurements of RQ. With minor modifications, this setup could be used in application to measure soil microbial respiratory behavior as oxygen is used up and conditions begin shifting to anaerobic, in effort to better understand the sensitivity and contribution of anaerobic respiration to bulk soil respiration.
Our thoughts are a single CUE that would best represent this period of glucose driven respiration would be a respiration weighted average of CUE over the period when RQ is ≥ 1. We believe this method is the most appropriate representation of CUE of the soil microbes specific to the period of time when glucose is the primary substrate.
We acknowledge that only considering C, H, O, and N is indeed a simplification of the real ecosystem and we hope that this work serves as a stepping stone and proof of concept. However, we are assured from Figure 5, that this fairly succinct simplification does effectively capture the process of biosynthesis in a manner which is approachable and compares very favorably with a carbon budget. However, further work could be done to provide an even more descriptive reaction to derive a more complete stoichiometric representation of soil microbes. However, it is also important to remember that other work has shown that microbial stoichiometry can vary with environmental conditions, so a more precise stoichiometry may be more applicable in certain use cases, but not all. Additional elements that could be included in this are phosphorus or sulfur, however their stoichiometric contributions are quite small in comparison to C, H, O, and N.
Comments Addressed:
Multiple lines : fixed in text citations.
Line 110, 298: reformatted Equations.
Lines 113 - 139: Moved appendix reaction derivation and added more derivation steps for readers to follow.
Lines 167 - 169 : Added sentence justifying our choice to add glucose as solid fine powder.
Lines 257, 258, 267-269: Added in figure numbers.
Line 329 : Added in reason for showing one replicate and method of choosing replicate, and why.
Lines 428 - 439: Added statements about how we think a prescribed CUE for the soils should be calculated and discussed the limitation of only using CHON elements to represent the system, as well as which amendment sizes are recommended for further experiments and future applications of this research.
Changes have been made to the working document and will be included in the next upload.
Citation: https://doi.org/10.5194/egusphere-2024-1757-AC2
-
AC2: 'Reply on RC2', Kyle Smart, 21 Jul 2024
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