Potential for synergy in soil inoculation for nature restoration by mixing inocula from different successional stages

Background and aims

Due to the well-known importance of biocrusts for several ecosystem properties linked to soil functionality, we aim to go deeper into the physiological performance of biocrusts components. Possible functional convergences in the physiology of biocrust constituents would facilitate the understanding of both species and genus distributional patterns and improve the possibility of modelling their response to climate change.

Methods

We measured gas exchange in the laboratory under controlled conditions of lichen- and moss-dominated biocrusts from four environmentally different locations in Europe. Field data were used to determine the natural hydration sources that drive metabolic activity of biocrusts.

Results

Our results show different activity drivers at the four sites. Within site analyses showed similar C fixation for the different crust types in the three sites without hydric stress whilst light use related parameters and respiration at 15 °C were similar in the between sites analyses. There were significant differences in water relations between the biocrusts types, with moss-dominated crusts showing higher maximum and optimum water contents.

Conclusions

The functional type approach for biocrusts can be justified from a physiological perspective when similar values are found in the within and between site analyses, the latter indicating habitat independent adaptation patterns. Our multi-site analyses for biocrusts functional performance provide comparisons of C fluxes and water relations in the plant-soil interface that will help to understand the adaptation ability of these communities to possible environmental changes.

     

What stabilizes biological soil crusts in the Negev Desert?

Aims

Biological soil crusts (biocrusts) are widespread in many drylands, where plant growth is limited due to water scarcity. One of their most important functions is the stabilization of the topsoil, particularly in regions with sandy soils prone to desertification. Since the mechanisms playing a role in soil stabilization are poorly understood, this study aims to shed light on the connection between crust stability and different cementing agents.

Methods

We measured the penetration resistance and the concentrations of different cementing agents of biocrusts in the Israeli Negev Desert. Structural equation modelling was performed to examine the direct and indirect effects of the variables analyzed and identify variables that are best able to explain the observed patterns of penetration resistance.

Results

All observed variables showed a high variability within and between sites. Structural equation modelling revealed that the main parameters explaining penetration resistance are the content of fines and the electrical conductivity, while carbonates and organic carbon only have an indirect effect.

Conclusions

Our results suggest that adding silt and clay to (natural or induced) biocrusts is very likely to produce stronger, more stable crusts, which will be more effective in combating desertification and improve their ability to survive trampling by livestock.

     

Trade-off between C and N recycling and N<Subscript>2</Subscript>O emissions of soils with summer cover crops in subtropical agrosystems

Aims

Biological soil crusts (biocrusts) are soil-surface communities in drylands, dominated by cyanobacteria, mosses, and lichens. They provide key ecosystem functions by increasing soil stability and influencing soil hydrologic, nutrient, and carbon cycles. Because of this, methods to reestablish biocrusts in damaged drylands are needed. Here we test the reintroduction of field-collected vs. greenhouse-cultured biocrusts for rehabilitation.

Methods

We collected biocrusts for 1) direct reapplication, and 2) artificial cultivation under varying hydration regimes. We added field-collected and cultivated biocrusts (with and without hardening treatments) to bare field plots and monitored establishment.

Results

Both field-collected and cultivated cyanobacteria increased cover dramatically during the experimental period. Cultivated biocrusts established more rapidly than field-collected biocrusts, attaining ~82% cover in only one year, but addition of field-collected biocrusts led to higher species richness, biomass (as assessed by chlorophyll a) and level of development. Mosses and lichens did not establish well in either case, but late successional cover was affected by hardening and culture conditions.

Conclusions

This study provides further evidence that it is possible to culture biocrust components from later successional materials and reestablish cultured organisms in the field. However, more research is needed into effective reclamation techniques.

     

Establishing rates of lateral expansion of cyanobacterial biological soil crusts for optimal restoration

Aims

Biocrusts that form on topsoils contribute ecosystem services to drylands, and their loss under anthropogenic pressure has negative ecological consequences. Therefore, development of biocrust inoculation technology for restoration is of interest. This requires knowledge of biocrust growth and dispersal. To contribute to this, we determined the speed at which biocrusts expand laterally based on the self-propelled motility of cyanobacteria.

Methodology

We inoculated sterile soil with natural biocrusts and incubated them over a year in a greenhouse under conditions mimicking local precipitation, monitoring the crust’s lateral expansion using time-course photography, chlorophyll a content, and microscopic inspection. Concurrent uninoculated controls served to monitor, and discount, natural inoculation by aeolian propagules.

Results

While the expansion front was highly variable in space, biocrusts expanded in the order of 2 cm month^?1, but only in seasons with moderate temperatures (Spring and Fall). Microcoleus vaginatus, Microcoleus steenstrupii, and Scytonema spp. advanced at averages of 1 cm month^?1, the crust advance front being preferentially driven by specialized propagules (hormogonia). These rates are within expectations based on instantaneous gliding motility speeds of cyanobacteria.

Conclusions

Based on the expansion capability of biocrusts during growth seasons, greenhouse inoculum units can be optimally spaced to fill 4–8 cm gaps.

     

Carbon mineralization and microbial activity in agricultural topsoil and subsoil as regulated by root nitrogen and recalcitrant carbon concentrations

Background and aims

Biocrust morphology is often used to infer ecological function, but morphologies vary widely in pigmentation and thickness. Little is known about the links between biocrust morphology and the composition of constituent microbial community. This study aimed to examine these links using dryland crusts varying in stage and morphology.

Methods

We compared the microbial composition of three biocrust developmental stages (Early, Mid, Late) with bare soil (Bare) using high Miseq Illumina sequencing. We used standard diversity measures and network analysis to explore how microbe-microbe associations changed with biocrust stage.

Results

Biocrust richness and diversity increased with increasing stage, and there were marked differences in the microbial signatures among stages. Bare and Late stages were dominated by Alphaproteobacteria, but Cyanobacteria was the dominant phylum in Early and Mid stages. The greatest differences in microbial taxa were between Bare and Late stages. Network analysis indicated highly-connected hubs indicative of small networks.

Conclusions

Our results indicate that readily discernible biocrust features may be good indicators of microbial composition and structure. These findings are important for land managers seeking to use biocrusts as indicators of ecosystem health and function. Treating biocrusts as a single unit without considering crust stage is likely to provide misleading information on their functional roles.

     

Fertilization,soil and plant community characteristics determine soil microbial activity in managed temperate grasslands

Background and aims

Contaminated soils can impede germination and growth of selected plant species, restricting effective phytoremediation strategies. The purpose of the present study was to enhance the germination and growth of saltgrass [Distichlis spicata (L.) Greene] by evaluating the efficacy of certain seed pretreatments and soil amendments.

Methods

Ten seed pretreatment methods, two amendments, three soil depths and five saline levels were tested under greenhouse conditions.

Results

Saltgrass germination and growth were negatively correlated with increasing salinity levels when NaCl > 85.6 mM. Among ten seed pretreatments (stratification + Proxy 24 h, hot water + Proxy 24 h, stratification, hot water + Proxy 48 h, Proxy 48 h, Proxy 24 h, hot water, scarification, gibberellins, and KMnO₄), the two best methods were stratification + Proxy 24 h and hot water + Proxy 24 h for enhancing saltgrass germination, with the latter pretreatment being especially useful because of its shorter preparation time and high germination rates. Proxy is a commercial ethephon product. Potting soil (5.0 cm depth) was found to be the best amendment for saltgrass germination and growth in hydrocarbon-contaminated soils.

Conclusion

We conclude that direct seeding of saline soils contaminated with petroleum hydrocarbons is a feasible phytoremediation strategy provided that appropriate seed pretreatments and amendments are utilized.

     

The effect of local climate and soil drainage on the environmental impact of grass-based milk production

Purpose

System modelling and life cycle assessment (LCA) were used to assess the climate change, acidification and eutrophication impacts of milk production using spring calving pasture-based system. The objective of the study was to evaluate the effect of climate and soil resources on the environmental impact per unit milk produced at the farm gate from low-cost, grass-based rotational-grazing dairy production.

Methods

A dairy system model, Dairy_sim, designed to identify optimum grass-based spring calving production systems considering the interaction between climate and soil resources was tested using the Irish National Dairy Blueprint and then used to assess regional differences of system management with well, moderately, mixed moderately-poorly and poorly drained soil resources available. Life cycle assessment was used to quantify environmental impacts of climate and soil drainage status. The Dairy_sim output was used as activity data for the LCA model.

Results and discussion

Differences were found in the management tactics influenced by climate and drainage resource. The impact of poor drainage reduced stocking rate, increased housing time and had greater need for later cut silage and more reliance on silage. Climate change, acidification and eutrophication impacts were greater for optimum management on poorly drained soil. The climate change ranged from 1.06 kg CO₂ eq./kg (well drained) to 1.18 kg CO₂ eq./kg (poorly drained) of energy corrected milk (ECM). The acidification and eutrophication ranged from 3.87 to 6.85 g SO₂ eq./kg ECM and 2.69 to 3.64 g PO₄ eq./kg ECM, respectively. Around 50% of poorly drained soil resource can be easily accommodated in dairy systems with little increase in environmental impact, where poor drained portion is utilised for silage.

Conclusions

LCA combined with a system optimization model revealed how dairy farm management practises constrained by poor land resource increased the environmental impact per unit product.

     

Effects of light intensity and quality on phycobiliprotein accumulation in the cyanobacterium <Emphasis Type="Italic">Nostoc sphaeroides</Emphasis> Kützing

Objectives

To assess the effects of light intensity and quality on the growth and phycobiliproteins (PBP) accumulation in Nostoc sphaeroides Kützing (N. sphaeroides).

Results

Dry weights, dry matter, protein, chlorophyll and PBP contents were higher under 90 μmol m^?2 s^?1 than under other intensities (both higher and lower). Phycocyanin and allophycocyanin increased with light intensity while phycoerythrin decreased. Fresh weights, protein and PBP contents increased at the highest rates under blue light. Red light resulted in higher values of dry matter, phycocyanin and chlorophyll a.

Conclusion

White light at 90 μmol m^?2 s^?1 or blue light 30 μmol m^?2 s^?1 were optimal for the growth and phycobiliprotein accumulation in N. sphaeroides.

     

A novel ethanol-tolerant laccase,Tvlac, from <Emphasis Type="Italic">Trametes versicolor</Emphasis>

Objectives

To produce and characterize novel laccases with ethanol tolerance from Trametes versicolor using agriculture by-products as energy source.

Results

Trametes versicolor 1017 produces two laccase isoenzymes with a total activity of 10 U ml^?1 within 8 days when using wheat bran and peanut powder as energy sources in liquid culture medium. A novel isoenzyme, named Tvlac, was identified, purified and characterized. Its optimum pH and temperature were from 4.5 to 5 and 55 to 60 °C, respectively. Its activity was stimulated by ethanol at 10 % (v/v) which increased the V ₀.

Conclusions

The biochemical properties of Tvlac substantiate the potential of this enzyme for applications under an aqueous ethanol mixture environment.

     

Ecofriendly demulsification of water in oil emulsions by an efficient biodemulsifier producing bacterium isolated from oil contaminated environment

Objective

Water in oil emulsions increase oil processing costs and cause damage to refinery equipment which necessitates demulsification. Since chemical demulsifiers cause environmental pollution, biodemulsifiers have been paid more attention. This study aims to identify biodemulsifier-producing bacteria from petroleum contaminated environments.

Results

As a result, several biodemulsifier producing strains were found that Stenotrophomonas sp. strain HS7 (accession number: MF445088) which produced a cell associated biodemulsifier showed the highest demulsifying ratio, 98.57% for water in kerosene and 66.28% for water in crude oil emulsion after 48 h. 35 °C, pH 7, 48 h incubation and ammonium nitrate as nitrogen source were optimum conditions for biodemulsifier production. Furthermore, it was found that hydrophobic carbon sources like as liquid paraffin is not preferred as the sole carbon source while a combination of various carbon sources including liquid paraffin will increase demulsification efficiency of the biodemulsifier.

Conclusions

The appropriate potential of this biodemulsifier strengthens the possibility of its application in industries especially petroleum industry.

     

Fallow associated with autumn-plough favors structure stability and storage of soil organic carbon compared to continuous maize cropping in Mollisols

Background and aims

Aggregate formation and stability of soil organic carbon (SOC) differ in different farming systems, probably due to differences in effects of tillage and residue management. This study used a 24-year field experiment to compare the effects of continuous maize cropping and natural fallow on aggregate formation and SOC storage in various aggregate-size classes and density fractions of a Chinese Mollisol.

Methods

Soils collected from the upper 0.2-m layer were wet-sieved into four aggregate-size classes (>2, 0.25–2, 0.053–0.25 and <0.053 mm) which were then fractionated into light, occluded and mineral C fractions. The concentrations of SOC and natural ¹³C abundance of each fraction in bulk soil and the aggregate classes were determined.

Results

Continuous maize cropping decreased the proportion of macro-aggregates (>0.25 mm) and increased that of micro-aggregates (<0.25 mm) compared to the initial value while the opposite was observed in the natural fallow system. The fallow system generally had greater SOC concentration in the occluded fraction, higher proportion of newly-derived C as % total SOC in the light fraction and greater contribution of total residue C to new C in macro-aggregates and light fractions compared to the continuous maize system. Furthermore, the fallow system resulted in shorter turnover time of SOC than the continuous maize system.

Conclusions

Natural fallow associated with autumn-plough improved soil structural stability and SOC storage while continuous maize cropping with residue removal decreased SOC sequestration and soil aggregate stability.

     

Does centrifugation matter? Centrifugal force and spinning time alter the plasma metabolome

Background

Centrifugation is an indispensable procedure for plasma sample preparation, but applied conditions can vary between labs.

Aim

Determine whether routinely used plasma centrifugation protocols (1500×g 10 min; 3000×g 5 min) influence non-targeted metabolomic analyses.

Methods

Nuclear magnetic resonance spectroscopy (NMR) and High Resolution Mass Spectrometry (HRMS) data were evaluated with sparse partial least squares discriminant analyses and compared with cell count measurements.

Results

Besides significant differences in platelet count, we identified substantial alterations in NMR and HRMS data related to the different centrifugation protocols.

Conclusion

Already minor differences in plasma centrifugation can significantly influence metabolomic patterns and potentially bias metabolomics studies.

     

Pig slurry acidification,separation technology and thermal conversion affect phosphorus availability in soil amended with the derived solid fractions,chars or ashes

Aims

The aim was to study effects of slurry acidification, separation technology and thermal processing on the availability of P in soil amended with the solid fraction of pig slurry.

Methods

Acidified and non-acidified slurry were separated using different technologies: screw press (SCR), decanting centrifuge (DEC) and drainage after chemical pretreatment (CHE). Solids and pyrolysed (400 °C or 600 °C, char) or combusted (625 °C, ash) solids were applied to two soils, with triple superphosphate (TSP) as a reference. Soil P availability was determined over 12 weeks using diffusive gradients in thin films (DGT).

Results

The initial availability of P from DEC and CHE solids was similar to TSP in both soils. After 6 weeks no significant difference was observed between DEC solids and DEC chars. Acidification did not significantly affect P availability in the solids-amended soils over time, but it did affect P availability with the thermally processed materials.

Conclusions

Application of separated pig slurry solids generally increased soil P availability initially, but declining with time, indicating soil P fixing reactions. Chars and ashes on the other hand showed lower initial P availability, but remained constant or increased slightly with chars yielding P availability similar to solids after 12 weeks.

     

Variation of soil respiration and its components in hemiboreal Norway spruce stands of different ages

Aims

Understanding the linkage of soil respiration (Rs) with forest development is essential for long-term C cycle models. We estimated the variation and temperature sensitivity (Q₁₀ value) of Rs and its hetero-, (Rh) and autotrophic (Ra) components in relation to abiotic and biotic factors in Norway spruce stands of different ages, and the effect of trenching on microbial and soil characteristics.

Methods

Trenching method was used to partition Rs into Rh and Ra. Ingrowth core method was used to estimate fine root production. Soil microbial biomass was measured using manometric respirometers.

Results

Rs varied in differently aged stands demonstrating non-linear response to development stage. The variation of Rs was explained by changes in biotic factors rather than by changes in soil microclimate. Rh was more sensitive to Ts than Rs or Ra. After 4 years of trenching soil pH, N, SOM and dehydrogenase activity were significantly changed in trenched plots compared to control plots.

Conclusions

Different Q₁₀ values of Rh and Ra in stands of different ages indicate the importance of Rs partitioning. Trenching should be used during a limited number of years because of the possible changes in chemical characteristics of soil and in the activity of soil microbial community.

     

A two-stage system coupling hydrolytic acidification with algal microcosms for treatment of wastewater from the manufacture of acrylonitrile butadiene styrene (ABS) resin

Objectives

To demonstrate the effectiveness of a novel two-stage system coupling hydrolytic acidification with algal microcosms for the treatment of acrylonitrile butadiene styrene (ABS) resin-manufacturing wastewater.

Results

After hydrolytic acidification, the BOD₅/COD ratio increased from 0.22 to 0.56, showing improved biodegradability of the wastewater. Coupled with hydrolytic acidification, the algal microcosms showed excellent capability of in-depth removal of COD, NH₃–N and phosphorus with removal rates 83, 100, and 89%, respectively, and aromatic pollutants, including benzene, were almost completely removed. The biomass concentration of Chlorella sp. increased from 5 × 10⁶ to 2.1 × 10⁷ cells/ml after wastewater treatment.

Conclusions

This two-stage coupling system achieved deep cleaning of the benzene-containing petrochemical wastewater while producing greater algae biomass resources at low cost.

     

Enhancement of cellulose degradation in freshwater sediments by a sediment microbial fuel cell

Objective

To demonstrate that an enhanced sediment microbial fuel cell (SMFC) system can accelerate the degradation of cellulose in fresh water sediments as the accumulation of cellulose in lake sediments may aggravate the lake marsh, increase organic matter content and result in rapid deterioration of water quality and damage the ecosystem.

Results

After 330 days the highest cellulose removal efficiency (72.7 ± 2.1 %) was achieved in the presence of a SMFC with a carbon nanotube decorated cathode, followed by a SMFC without the cathode decoration (64.4 ± 2.8 %). The lowest cellulose removal efficiency (47.9 ± 2.1 %) was in the absence of SMFC. The sediment characterization analysis confirmed that the carbon nanotube decorated cathode enhances the electron transfer rate in the SMFC and improves the dissolved organic matter oxidation rate.

Conclusion

This study offers a relatively simple and promising new method for cellulose degradation in sediment.

     

Effects of melatonin administration on embryo implantation and offspring growth in mice under different schedules of photoperiodic exposure

Background

Embryo implantation is crucial for animal reproduction. Unsuccessful embryo implantation leads to pregnancy failure, especially in human-assisted conception. Environmental factors have a profound impact on embryo implantation. Because people are being exposed to more light at night, the influence of long-term light exposure on embryo implantation should be explored.

Methods

The effects of long photoperiodic exposure and melatonin on embryo implantation and offspring growth were examined. Long photoperiodic exposure (18:6 h light:dark) was selected to resemble light pollution. Melatonin (10^?2, 10^?3, 10^?4, 10^?5 M) was added to the drinking water of mice starting at Day 1 (vaginal plugs) until delivery.

Results

Melatonin treatment (10^?4,10^?5 M) significantly increased litter sizes compared to untreated controls (12.9 ± 0.40 and 12.2 ± 1.01 vs. 11.5 ± 0.43; P < 0.05). The most effective concentration of melatonin (10^?4 M) was selected for further investigation. No remarkable differences were found between melatonin-treated mice and controls in terms of the pups’ birth weights, weaning survival rates, and weaning weights. Long photoperiodic exposure significantly reduced the number of implantation sites in treated mice compared to controls (light/dark, 12/12 h), and melatonin rescued this negative effect. Mechanistic studies revealed that melatonin enhanced the serum 17β-estradiol (E₂) levels in the pregnant mice and upregulated the expression of the receptors MT1 and MT2 and p53 in uterine tissue. All of these factors may contribute to the beneficial effects of melatonin on embryo implantation in mice.

Conclusion

Melatonin treatment was associated with beneficial effects in pregnant mice, especially those subjected to long photoperiodic exposure. This was achieved by enhanced embryo implantation. At the molecular level, melatonin administration probably increases the E₂ level during pregnancy and upregulates p53 expression by activating MT1/2 in the uterus. All of the changes may improve the microenvironment of the uterus and, thus, the outcomes of pregnancy.

     

Biocrusts enhance soil fertility and <Emphasis Type="Italic">Bromus tectorum</Emphasis> growth,and interact with warming to influence germination

Background and aims

Biocrusts are communities of cyanobacteria, mosses, and/or lichens found in drylands worldwide. Biocrusts are proposed to enhance soil fertility and productivity, but simultaneously act as a barrier to the invasive grass, Bromus tectorum, in western North America. Both biocrusts and B. tectorum are sensitive to climate change drivers, yet how their responses might interact to affect dryland ecosystems is unclear.

Methods

Using mesocosms with bare soil versus biocrust cover, we germinated B. tectorum seeds collected from warmed, warmed + watered, and ambient temperature plots within a long-term climate change experiment on the Colorado Plateau, USA. We characterized biocrust influences on soil fertility and grass germination, morphology, and chemistry.

Results

Biocrusts increased soil fertility and B. tectorum biomass, specific leaf area (SLA), and root:shoot ratios. Germination rates were unaffected by mesocosm cover-type. Biocrusts delayed germination timing while also interacting with the warmed treatment to advance, and with the warmed + watered treatment to delay germination.

Conclusions

Biocrusts promoted B. tectorum growth, likely through positive influence on soil fertility which was elevated in biocrust mesocosms, and interacted with seed treatment-provenance to affect germination. Understanding how anticipated losses of biocrusts will affect invasion dynamics will require further investigation of how plant plasticity/adaptation to specific climate drivers interact with soil and biocrust properties.

     

Mid- and long-term correlations of plasma metabolite concentrations measured by a targeted metabolomics approach

Introduction

Plasma metabolites measured by metabolomics techniques have been shown to be associated with chronic disease risk in epidemiological studies. However, in most prospective studies metabolomic profiles can only be obtained at a single time point and data on intra-individual variation in metabolite levels over time are sparse.

Objectives

Here, we evaluated the intra-individual variation in the concentrations of 177 metabolites over time using repeat blood samples of 104 adults (50% female).

Methods

Blood samples were obtained at a baseline visit between 1994 and 1998 and during two further examinations 14 and 15 years later. Plasma metabolite levels were quantified by tandem mass spectrometry with the MetaDisIDQ? p180 Kit. Intra-individual variation was assessed by Spearman’s correlation coefficients (ρ).

Results

Mid-term correlations over 1 year were good (ρ ≥ 0.7) for 5.1% and reasonable (ρ ≥ 0.4 < 0.7) for 61.0% of the metabolites, while long-term correlations over 15 years were good for 2.8% and reasonable for 27.1%. The strongest mid-term correlations between metabolite concentrations were observed for the acylcarnitine C₃–OH (0.72) and metabolites from the amino acid/biogenic amine groups, i.e. creatinine (0.83), proline (0.79), lysine (0.77), isoleucine (0.76), and ornithine (0.74). C₃–OH (0.78) as well as several amino acids/biogenic amines, i.e. ornithine (0.76), sarcosine (0.76), lysine (0.75), spermine (0.73), and glutamine (0.69) showed the strongest long-term correlations.

Conclusions

The biological reproducibility of plasma concentrations of a majority of metabolites occurs reasonable over 1 year. In contrast, concentrations of many metabolites seem to be affected by substantial intra-individual variation over 15 years.

     

Age dependent accumulation patterns of advanced glycation end product receptor (RAGE) ligands and binding intensities between RAGE and its ligands differ in the liver,kidney, and skeletal muscle

Background

Much evidence indicates receptor for advanced glycation end products (RAGE) related inflammation play essential roles during aging. However, the majority of studies have focused on advanced glycation end products (AGEs) and not on other RAGE ligands. In the present study, the authors evaluated whether the accumulation of RAGE ligands and binding intensities between RAGE and its ligands differ in kidney, liver, and skeletal muscle during aging.

Results

In C57BL/6 N mice aged 12 weeks, 12 months, and 22 months, ligands accumulation, binding intensities between RAGE and its ligands, activated macrophage infiltration, M1/M2 macrophage expression, glyoxalase-1expression, and signal pathways related to inflammation were evaluated. The RAGE ligands age-associated accumulation patterns were found to be organ dependent. Binding intensities between RAGE and its ligands in kidney and liver increased with age, but those in skeletal muscle were unchanged. Infiltration of activated macrophages in kidney and liver increased with age, but infiltration in the skeletal muscle was unchanged. M1 expression increased and M2 and glyoxalase-1 expression decreased with age in kidney and liver, but their expressions in skeletal muscle were not changed.

Conclusion

These findings indicate patterns of RAGE ligands accumulation, RAGE/ligands binding intensities, or inflammation markers changes during aging are organs dependent.