Effects of Waterlogging on Water Relations of Actively-growing and Dormant Sitka Spruce Seedlings

Two-year-old Sitka spruce [Picea sitchensis (Bong.) Carr.] seedlings,either actively growing or dormant, were waterlogged in a growthroom at 15 °C. Shoot and root growth, transpiration andleaf water potential were observed. In actively-growing plants shoot extension continued after waterlogging,though at a reduced rate, and shoots of dormant plants brokebud and extended during the waterlogging period. Root growthwas suppressed by waterlogging in both types of plant. The 22day waterlogging treatment eventually killed the actively-growingplants but plants which were dormant at the time of waterloggingwere more tolerant. Changes in plant water relations after waterloggingwere entirely different depending on the condition of the plantswhen the soil was flooded. Dormant plants showed a gradual reductionin transpiration and increased water stress over the waterloggingperiod; after the soil was drained leaf water potential increasedto equal the value of control plants which had been maintainedin a freely drained condition, but transpiration did not increaseuntil root growth began. Actively-growing plants exhibited amore complex behaviour, characterized by a very rapid reductionin transpiration after waterlogging, accompanied by a briefperiod of water stress, followed by a period of increasing transpirationrate in the absence of water stress. Finally a second reductionin transpiration occurred and water stress increased as theseedlings died. The importance of the stage of activity of theroot system to the response of plants to waterlogging is discussed. Picea sitchensis (Bong.) Carr., Sitka spruce, waterlogging, water relations, dormancy, transpiration, water potential     

Mechanisms for the increase in phosphorus uptake of waterlogged plants: soil phosphorus availability, root morphology and uptake kinetics

Waterlogging frequently reduces plant biomass allocation to roots. This response may result in a variety of alterations in mineral nutrition, which range from a proportional lowering of whole-plant nutrient concentration as a result of unchanged uptake per unit of root biomass, to a maintenance of nutrient concentration by means of an increase in uptake per unit of root biomass. The first objective of this paper was to test these two alternative hypothetical responses. In a pot experiment, we evaluated how plant P concentration of Paspalum dilatatum, (a waterlogging-tolerant grass from the Flooding Pampa, Argentina) was affected by waterlogging and P supply and how this related to changes in root-shoot ratio. Under both soil P levels waterlogging reduced root-shoot ratios, but did not reduce P concentration. Thus, uptake of P per unit of root biomass increased under waterlogging. Our second objective was to test three non-exclusive hypotheses about potential mechanisms for this increase in P uptake. We hypothesized that the greater P uptake per unit of root biomass was a consequence of: (1) an increase in soil P availability induced by waterlogging; (2) a change in root morphology, and/or (3) an increase in the intrinsic uptake capacity of each unit of root biomass. To test these hypotheses we evaluated (1) changes in P availability induced by waterlogging; (2) specific root length of waterlogged and control plants, and (3) P uptake kinetics in excised roots from waterlogged and control plants. The results supported the three hypotheses. Soil P avail-ability was higher during waterlogging periods, roots of waterlogged plants showed a morphology more favorable to nutrient uptake (finer roots) and these roots showed a higher physiological capacity to absorb P. The results suggest that both soil and plant mechanisms contributed to compensate, in terms of P nutrition, for the reduction in allocation to root growth. The rapid transformation of the P uptake system is likely an advantage for plants inhabiting frequently flooded environments with low P fertility, like the Flooding Pampa. This advantage would be one of the reasons for the increased relative abundance of P. dilatatum in the community after waterlogging periods. Received: 15 February 1997 / Accepted: 20 May 1997     

The Induction of Root Dormancy in Picea sitchensis (Bong.) Carr. by Abscisic Acid

Abscisic acid at 10⁴ M inhibited elongation and induced dormancyin roots of Sitka spruce cuttings growing in solution culture.The dormancy was characterized by superficial browning whichprogressed towards the root tip and by the development of alayer of lignified and suberized cells around the root apex.The dormant roots remained alive and resumed growth on transferto ABA-free solutions. The induction of dormancy was confinedto those roots actually bathed in the ABA solution and a rangeof conditions which affected shoot activity did not alter theresponse. By contrast, the time for which the roots had beenelongating strongly influenced root dormancy since it was notinduced in roots which had recently begun to elongate in solutionculture. The possible role of ABA in inducing root dormancyunder adverse conditions is discussed.     

Plant growth, nutrient acquisition and mycorrhizal symbioses of a waterlogging tolerant legume (Lotus glaber Mill.) in a saline-sodic soil

Seedlings of Lotus glaberMill., were grown in a native saline-sodic soil in a greenhouse for 50 days and then subjected to waterlogging for an additional period of 40 days. The effect of soil waterlogging was evaluated by measuring plant growth allocation, mineral nutrition and soil chemical properties. Rhizobiumnodules and mycorrhizal colonisation in L. glaberroots were measured before and after waterlogging. Compared to control plants, waterlogged plants had decreased root/shoot ratio, lower number of stems per plant, lower specific root length and less allocation of P and N to roots. Waterlogged plants showed increased N and P concentrations in plant tissues, larger root crown diameter and longer internodes. Available N and P and organic P, pH and amorphous iron increased in waterlogged soil, but total N, EC and exchangeable sodium were not changed. Soil waterlogging decreased root length colonised by arbuscular mycorrhizal (AM) fungi, arbuscular colonisation and number of entry points per unit of root length colonised. Waterlogging also increased vesicle colonisation and Rhizobium nodules on roots. AM fungal spore density was lower at the end of the experiment in non-waterlogged soil but was not reduced under waterlogging. The results indicate that L. glaber can grow, become nodulated by Rhizobium and colonised by mycorrhizas under waterlogged condition. The responses of L. glaber may be related its ability to form aerenchyma.     

Fine root biomass in two black spruce stands in interior Alaska: effects of different permafrost conditions

Key message

In black spruce stands on permafrost, trees and understory plants showed higher biomass allocation especially to ‘thin’ fine roots (diam. < 0.5 mm) when growing on shallower permafrost table.

Abstract

Black spruce (Picea mariana) forests in interior Alaska are located on permafrost and show greater below-ground biomass allocation than non-permafrost forests. However, information on fine roots (roots <2 mm in diameter), which have a key role in nutrient uptake and below-ground carbon flux, is still limited especially for effects of different permafrost conditions. In this study, we examined fine root biomass in two black spruce stands with different depths to the permafrost table. In the shallow permafrost (SP) plot, fine root biomass of black spruce trees was 70 % of that in the deep permafrost (DP) plot. In contrast, ratio of the fine root biomass to above-ground biomass was greater in the SP plot than in the DP plot. Understory plants, on the other hand, showed larger fine root biomass in the SP plot than in the DP plot, whereas their above-ground biomass was similar between the two plots. In addition, biomass proportion of ‘thin’ fine roots (diam. <0.5 mm) in total fine roots was greater in the SP plot than in the DP plot. These results suggest that black spruce trees and understory plants could increase biomass allocation to fine roots for efficient below-ground resource acquisition from colder environments with shallower permafrost table. In the SP plot, fine roots of understory plants accounted for 30 % of the stand fine root biomass, suggesting that understory plants such as Ledum and Vaccinium spp. would have significant contribution to below-ground carbon dynamics in permafrost forests.

     

The Influence of Pine on the Form of Sitka Spruce Fine Roots

The form of fine roots of Sitka spruce (Picea sitchensis Bong.Carr.) when grown in immediate proximity to roots of Scots pine(Pinus sylvestris L.) and when grown separately was comparedusing split-root systems. When spruce roots were intimatelymixed with pine roots the mean length of individual spruce lateralswas significantly greater, while the length: weight ratio andnumber of root tips: weight ratio were smaller than when grownalone. A similar alteration in growth strategy was achievedby direct addition of mineral nitrogen. Key words: Fine root form, pine-spruce interactions     

Aerenchyma formation and radial O2 loss along adventitious roots of wheat with only the apical root portion exposed to O2 deficiency

This study investigated aerenchyma formation and function in adventitious roots of wheat (Triticum aestivum L.) when only a part of the root system was exposed to O₂ deficiency. Two experimental systems were used: (1) plants in soil waterlogged at 200 mm below the surface; or (2) a nutrient solution system with only the apical region of a single root exposed to deoxygenated stagnant agar solution with the remainder of the root system in aerated nutrient solution. Porosity increased two‐ to three‐fold along the entire length of the adventitious roots that grew into the water‐saturated zone 200 mm below the soil surface, and also increased in roots that grew in the aerobic soil above the water‐saturated zone. Likewise, adventitious roots with only the tips growing into deoxygenated stagnant agar solution developed aerenchyma along the entire main axis. Measurements of radial O₂ loss (ROL), taken using root‐sleeving O₂ electrodes, showed this aerenchyma was functional in conducting O_2. The ROL measured near tips of intact roots in deoxygenated stagnant agar solution, while the basal part of the root remained in aerated solution, was sustained when the atmosphere around the shoot was replaced by N₂. This illustrates the importance of O₂ diffusion into the basal regions of roots within an aerobic zone, and the subsequent longitudinal movement of O₂ within the aerenchyma, to supply O₂ to the tip growing in an O₂ deficient zone.     

Constraints of root response to waterlogging in Alisma triviale

To understand the economics of root aerenchyma formation in wetland plants, we investigated in detail the response of Alisma triviale to waterlogging. We hypothesized costs being associated with development of a large root air space. In three out-door pot experiments, seedlings (1 experiment) and mature plants (2 experiments) were grown under waterlogged and drained conditions for up to 2?months. Waterlogging promoted growth, and was associated with increased root porosity and decreased root density (fresh mass per volume). The increased formation of aerenchyma was associated with a higher root dry matter content for a given root density. Despite improved growth and earlier flowering, the waterlogged plants also showed signs of being constrained by the anoxic substrate, such as shallower roots, and a higher leaf dry matter content. The formation of aerenchyma was associated with costs, such as increased root dry matter content and reduced metaxylem vessel diameter. The faster growth of the seedlings under the waterlogged conditions, despite some signs of being stressed, was possibly a result of decreased requirements to allocate biomass below ground. In mature plants the increased aerenchyma allowed deeper root penetration, and ameliorated the effects of anoxia, reducing the differences in plant traits between the treatments.     

Mechanisms of waterlogging tolerance in wheat – a review of root and shoot physiology

We review the detrimental effects of waterlogging on physiology, growth and yield of wheat. We highlight traits contributing to waterlogging tolerance and genetic diversity in wheat. Death of seminal roots and restriction of adventitious root length due to O₂ deficiency result in low root:shoot ratio. Genotypes differ in seminal root anoxia tolerance, but mechanisms remain to be established; ethanol production rates do not explain anoxia tolerance. Root tip survival is short‐term, and thereafter, seminal root re‐growth upon re‐aeration is limited. Genotypes differ in adventitious root numbers and in aerenchyma formation within these roots, resulting in varying waterlogging tolerances. Root extension is restricted by capacity for internal O₂ movement to the apex. Sub‐optimal O₂ restricts root N uptake and translocation to the shoots, with N deficiency causing reduced shoot growth and grain yield. Although photosynthesis declines, sugars typically accumulate in shoots of waterlogged plants. Mn or Fe toxicity might occur in shoots of wheat on strongly acidic soils, but probably not more widely. Future breeding for waterlogging tolerance should focus on root internal aeration and better N‐use efficiency; exploiting the genetic diversity in wheat for these and other traits should enable improvement of waterlogging tolerance.     

Control of Water Loss by Actively Growing Sitka Spruce Seedlings after Transplanting

The roots of actively growing Sitka spruce seedlings were disturbedand damaged to different degrees by transplanting treatmentscarried out in a growthroom. The treatments stopped root extensionand after 2–4 h the transpiration rate of treated plantsdeclined, indicating closure of the stomata, and transpirationcontinued to decrease for a few days. The hydraulic conductivityof the root system was reduced slightly by the treatment butleaf water potential increased or remained unchanged, thereforeno water stress occurred to account for closure of the stomata.Measurements on detached shoots indicated that the water storagecapacity was too small to accommodate changes in the vascularwater supply without reducing leaf water potential. The measuredresponses of the plant were not affected by girdling the baseof the stem prior to root treatment. It is suggested that thetranspiration rate of treated plants was reduced as a resultof a chemical stimulus originating in the root and acting onthe stomata.     

Roots of willow (Salix viminalis L.) show marked tolerance to oxygen shortage in flooded soils and in solution culture

Responses to soil flooding and oxygen shortage were studied in field, glasshouse and controlled environment conditions. Established stools ofSalix viminalis L., were compared at five field sites in close proximity but with contrasting water table levels and flooding intensities during the preceding winter. There was no marked effect of site on shoot extension rate, time to half maximum length or final length attained. When rooted cuttings were waterlogged for 4 weeks in a glasshouse, soil redox potentials quickly decreased to below zero. Shoot extension was slowed after a delay of 20 d, while, in the upper 100 mm of soil, formation and outgrowth of unbranched adventitious roots with enhanced aerenchyma development was promoted after 7 d. At depths of 100–200 mm and 200–300 mm, extension by existing root axes was halted by soil flooding, while adventitious roots from above failed to penetrate these deeper zones. After 4 weeks waterlogging, all arrested root tips recommenced elongation when the soil was drained; their extension rates exceeding those of roots that were well-drained throughout. Growth in fresh mass was also stimulated. The additional aerenchyma found in adventitious roots in the upper 100 mm of soil may have been ethylene regulated since gas space development was inhibited by silver nitrate, an ethylene action inhibitor. The effectiveness of aerenchyma was tested by blocking the entry of atmospheric oxygen into plants with lanolin applied to lenticels of woody shoots of plants grown in solution culture. Root extension was halved, while shoot growth remained unaffected. H Lambers Section editor     

The arbuscular mycorrhizal fungus Diversispora spurca ameliorates effects of waterlogging on growth,root system architecture and antioxidant enzyme activities of citrus seedlings

Citrus plants are often exposed to heavy rain and subsequent periods of soil waterlogging which severely restrict tree growth. We assessed the effect of one arbuscular mycorrhizal fungus species (Diversispora spurca) on growth, root system architecture (RSA), and antioxidant enzyme activities of young citrus (Citrus junos) seedlings. Waterlogging for 37 d significantly restricted mycorrhizal colonization but increased the number of entry points and vesicles. Compared with non-mycorrhizal controls, mycorrhizal seedlings had significantly greater plant height, fresh mass, total root and taproot lengths, projected and surface root areas, root volume, and numbers of lst, 2nd and 3rd order lateral roots regardless of waterlogging treatment. D. spurca significantly increased root catalase (CAT) activity in non-stressed seedlings and increased root soluble protein concentration and leaf CAT activity in waterlogged seedlings, thereby inducing lower oxidative damage. These results suggest that D. spurca ameliorates effects of waterlogging on growth, RSA and antioxidant enzyme activities.     

Aerenchyma and an Inducible Barrier to Radial Oxygen Loss Facilitate Root Aeration in Upland, Paddy and Deep-water Rice (Oryza sativa L.)

The present study evaluated waterlogging tolerance, root porosityand radial O₂ loss (ROL) from the adventitious roots, of sevenupland, three paddy, and two deep-water genotypes of rice (Oryzasativa L.). Upland types, with the exception of one genotype,were as tolerant of 30 d soil waterlogging as the paddyand deep-water types. In all but one of the 12 genotypes, thenumber of adventitious roots per stem increased for plants grownin waterlogged, compared with drained, soil. When grown in stagnantdeoxygenated nutrient solution, genotypic variation was evidentfor root porosity and rates of ROL, but there was no overalldifference between plants from the three cultural types. Adventitiousroot porosity increased from 20–26 % for plants grownin aerated solution to 29–41 % for plants grown instagnant solution. Growth in stagnant solution also induceda ‘tight’ barrier to ROL in the basal regions ofadventitious roots of five of the seven upland types, all threepaddy types, and the two deep-water types. The enhanced porosityprovided a low resistance pathway for O₂ movement to the roottip, and the barrier to ROL in basal zones would have furtherenhanced longitudinal O₂ diffusion towards the apex, by diminishinglosses to the rhizosphere. The plasticity in root physiology,as described above, presumably contributes to the ability ofrice to grow in diverse environments that differ markedly insoil waterlogging, such as drained upland soils as well as waterloggedpaddy fields.     

Spatial variation in root system activity of tomato (Solanum lycopersicum L.) in response to short and long-term waterlogging as determined by 15N uptake

Background and aim

Root system activity is affected by abiotic stresses, which often creates spatial differences in root conditions. This is expected to influence plants ability to cope with suboptimal conditions.

Methods

Changes in root system activity were determined as ¹⁵N root uptake in top and bottom layers of potted tomato plants (Solanum lycopersicum L.), after waterlogging the bottom layer for 24 h or 5 d. The plants were grown in peat-based media; non-compacted or highly-compacted, resulting in differences in gas diffusion, air permeability and oxygen availability.

Results

The roots were affected by short-term waterlogging (24 h) by decreasing uptake in the bottom layer and increasing uptake in the pot top layer. Long-term waterlogging (5 d) decreased the ¹⁵N root uptake more in both layers. Root uptake recovered fast (within 6 h) after short-term waterlogging, whereas recovery of long-term waterlogged roots took more than 24 h, suggesting production of new root biomass. Despite affecting physical properties, medium compaction did not affect root uptake. Aboveground biomass was affected by waterlogging by increasing the dry matter percentage, decreasing nitrogen (N) percentage and increasing starch content.

Conclusions

The results confirmed that root uptake in different layers of small pots could be distinguished by the ¹⁵N technique, which was applicable under potentially denitrifying conditions. The results demonstrated that during short-term stress in part of the root system plants increased uptake from the non-affected parts of the root system, probably as compensation for suboptimal conditions.     

Nitrogen status of cotton subjected to two short term periods of waterlogging of varying severity using a sloping plot water-table facility

Summary Cotton is reported to be susceptible to waterlogging, and there is evidence that some of the symptoms shown by waterlogged plants are due to impaired uptake of nitrogen. To investigate this for cotton, the nitrogen nutrition of a field-grown crop was monitored when the plants were subjected to two short term periods of waterlogging of varying severity using a sloping plot water-table facility. Growth of severely waterlogged cotton decreased after 4 days in the first and second floodings, and these plants were wilted by the end of the first flooding but not the second. Waterlogging resulted in decreased concentrations of total-N and especially NO ₃ ^– –N in the petiole and lamina of the youngest fully-expanded leaf. Uptake of N by waterlogged plants occurred, but was not as great as for well-aerated plants. The nitrate reductase activity of leaves was much lower in waterlogged plants. Stumps of detopped waterlogged plants did not exude sylem sap at the end of the first flooding, suggesting impaired solute uptake due to damaged roots. However, xylem exudate was obtained from stumps of waterlogged plants at the end of the second flooding, indicating adaptive changes to the root systems of these plants. Although cotton is reported to reduce little NO ₃ ^– –N in its roots, analysis of xylem exudate showed that about half of the N exported by roots was as amino compounds. The concentration of amino compounds in xylem exudate from severely waterlogged plants was higher than in well-aerated plants. It was concluded that the growth reduction in waterlogged cotton was due partly to induced N-deficiency.     

Relationship between mineral coating on roots and yield performance of wheat under waterlogging stress

Waterlogging stress limits the productivity of winter wheat(Triticum aestivum L. emend. Thell.) in many parts of the world.When wheat is grown under waterlogged conditions, a reddish-brownmineral coating can form on the epidermal surface of the roots.In wetland plants such as rice, the amount of mineral coatingformed on root surfaces is positively related to yield. Thisstudy was conducted to determine whether mineral coating onthe roots of wheat is related to yield potential under waterloggedconditions. Root mineral coatings formed under waterlogged conditionswere studied in 12 cultivars and two breeding lines over threeyears of greenhouse pot studies. Soil redox potential in thewaterlogged treatment ranged between - 46 and 171 mV, and grainyield was suppressed by 28–49% compared to well-drainedcontrols. Mineral coating formed on the roots from the waterloggedtreatment was determined to be composed primarily of iron, basedon ICP elemental analysis, iron-specific staining, and ion-mappingby scanning electron microscopy using an X-ray detector. Of11 elements quantified by ICP spectroscopy, six were significantlyaffected by waterlogging treatment, and three of these, Fe,Mn and P, were well-correlated negatively to yield. Aerenchymaformation in the heavily coated waterlogged roots appeared todisrupt the internal root structure, and exceeded 40% of cross-sectionalarea in one cultivar. Unlike rice, which shows a positive relationshipbetween oxygen release from roots, grain yield and mineral coating,in winter wheat, the amount of mineral coating is negativelyrelated to grain yield under waterlogged conditions. Key words: Wheat, mineral coating, waterlogging, aerenchyma, hypoxia     

The response of sorghum and sunflower to short-term waterlogging

Summary Sorghum and sunflower were waterlogged for nine days during the vegetative, floral initiation/buds-visible or anthesis stage of growth under glasshouse conditions to observe the effects on root growth and development. In addition, some plants were waterlogged at all three stages to observe any adaptations induced by waterlogging. The most marked effects occurred at the initiation/buds-visible stage where a 30% reduction in root length and a 40% reduction in root dry weight of sorghum occurred with comparable figures for sunflower being 50 and 60% respectively. Generally, sorghum roots had a higher porosity than sunflower which may contribute to its greater tolerance to waterlogging. The observed changes in root growth are discussed in relation to previously documented effects of waterlogging on growth of the two species and changes which occur in the soil environment.     

Specific root area: A soil characteristic

The root systems of all plants adapt to soil conditions. One of the ways Norway spruce adapts is a regulation of absorbing root surface area, which can be viewed as an ecomorphological index of soil conditions. Minimal specific root surface area of absorbing roots of Norway spruce (28–29 m²kg⁻¹) responds to optimal soil conditions. This was achieved at a soil bulk density of about 1.2 gcm⁻³.     

Distribution of roots and root nodules and biomass allocation in young intensively managed grey alder stands on a peat bog

Development of below-ground biomass and biomass allocation were studied in two different stands of young grey alder stands growing on a peat bog. Both stands were given the same fertilization and irrigation treatment. The roots were investigated from 1) open plastic tubes enclosing the complete root systems in 1982, and 2) root cores 1984–86. Coarse roots (diameter>1 mm) were mainly found close to the trunk of the trees while fine roots (≤1 mm) were more evenly distributed in the stands. Root nodules were intermediate in distribution. The root systems were shallow, with more than 90% of the biomass in the uppermost 9–10 cm of the soil, probably because of low oxygen availability in the peat soil. The biomass allocation to the above-ground parts increased during the study period.     

Effects of waterlogging on soil aeration and on root and shoot growth and yield of winter oats (Avena sativa L.)

Summary Winter oats were grown outdoors in lysimeters containing monoliths of a sandy loam soil. The soil was either freely-drained throughout the experiment or waterlogged to the soil surface from mid-January until mid-April. After the start of waterlogging the oxygen flux density decreased most rapidly nearer the soil surface and in the upper 50 cm declined to zero. At 80 cm depth the oxygen flux density at the end of the waterlogging still had not diminished to zero. While the soil was waterlogged root growth was negligible in the 20–50 cm zone of the soil profile, whereas below that depth root growth continued, reaching 95 cm by the end of the treatment. During the latter part of the waterlogging period root growth resumed in the upper 10 cm, and in the upper 2.5 cm was greater than in the freelydrained treatment.At the end of the waterlogging period, the total root length and shoot dry weights were 77 and 60% of those in the freely-drained treatment, tillering was restricted and leaf area index diminished. However, by anthesis, root length and shoot weights of the plants that had been waterlogged were only 10 and 12% less respectively than for the freely-drained plants. At harvest, total dry matter and grain yields were only 9% less, the latter largely through fewer grains per panicle.