Using branch and basal trunk sap flow measurements to estimate whole-plant water capacitance: a caution

Thermometric sap flow sensors are widely used to measure water flow in roots, stems and branches of plants. Comparison of the timing of flow in branches and stems has been used to estimate water capacitance of large trees. We review studies of sap flow in branches and present our own data to show that there is wide variation in the patterns and timing of sap flow of branches in different parts of the crown, owing to the course of daily solar illuminance. In contiguous forest, east-facing and upper branches are illuminated earlier than west-facing and lower branches and most capacitance studies do not include adequate information about branch sampling regimes relative to the overall pattern of crown illuminance, raising questions about the accuracy of capacitance estimates. Measuring only upper branches and normalising these results to represent the entire crown is dangerous because flows at the stem base likely peak in response to maximum crown illuminance (and transpiration) and this will differ compared to the timing of peak flows in upper branches. We suggest that the magnitude of flow lags between branches and stems needs further study, with careful attention to branch position and method application before a robust understanding of capacitance, particularly in woody tissues of large trees, can be formed. We did not detect flow lags in the world’s tallest and largest tree species Sequoia sempervirens and Sequoiadendron giganteum, despite measurement along large pathlengths (∼57 and 85 m), which raises questions as to why large flow lags are often recorded for much smaller species. One conspicuous possibility is the different methods used among studies. Constant-heating methods such as the thermal dissipation probe (and also heat balance methods) include heat capacitance behaviour due to warming of wood tissues, which delays the response of the sensors to changing sap flow conditions. We argue that methods with intrinsic heat-capacitance present dangers when trying to measure water-capacitance in trees. In this respect heat pulse methods hold an advantage.     

Time lags for xylem and stem diameter variations in a Scots pine tree

Diameter variations in the xylem and whole stem (i.e. over bark) stem of a Scots pine (Pinus sylvestris L.) tree were measured at four heights over a 23 d period at 5 min intervals. Cross‐correlation analysis was used to calculate time lags between the measurements. Xylem diameter measurements at the different heights had time lags varying from 10 to 50 min, measurements at the lower heights lagging behind the most. This result was in good agreement with the cohesion theory of transpiration. For the whole stem diameter measurements, the treetop lagged behind all other heights and the shortest lags were midway along the stem. Changes in whole stem diameter always lagged behind those of xylem stem diameter (30–110 min), and at all heights. The considerable differences in the behaviour of xylem and whole stem diameter support the Münch hypothesis of phloem flow. Time lags calculated separately for the shrinkage (morning) and swelling (afternoon) periods indicated shorter time lags during the swelling periods. The non‐destructive methods used show promise in the simultaneous study of flow dynamics of xylem and phloem in trees.     

Significance of the Occurrence of Time Lags in the Transmission of Hydraulic Shock Waves through Plant Stems

Hydraulic shock waves were introduced into plants by pruning.These shock waves were monitored by a sap flux meter attachedto the plant at some distance from the excision. The measuredshock waves occurred a short time after pruning. An electricalanalogy is proposed to explain the time lags. The analogy suggestedthat, in plants undergoing auto-oscillatory behaviour, thereshould be a time lag in the transmission of oscillations insap flux along a stem. A plant undergoing auto-oscillatory watermetabolism showed a time lag of this type.     

Time lags in ecological systems.

A model representing ten species in a four trophic level community is constructed by using Volterra's equations including time lags, and is solved numerically for some values of the parameters. Classical discrete and continuous time lags yield similar results; simulating with discrete time lags thus appears useful. Parasitic versus predatory structures are compared by measuring the amplitudes of oscillations and the time taken to settle down to equilibrium, and give the following qualitative conclusions for the model.An increase in the carrying capacity of a trophic level increases the destabilizing influence of time lag in that level, this increase is more marked in predatory structures. Time lags appear to cause more violent oscillations in species strongly linked to a predatory system than in weakly linked species. The oscillatory period increases in predatory systems as the time lag is moved to higher levels, while it decreases in parasitic systems.     

The Effect of Bud Position on Branch Growth and Bud Abscission in Quercus petraea (Matt.) Liebl.

The time at which a bud began to expand was related to its positionnot only on an individual shoot but also within the crown. Thedistribution of buds and branches on the shoot was uneven; theshoot tip, where they were densely clustered, was termed the‘whorl; and the remainder of the shoot, where they werewidely spaced, the ‘interwhorl’ stem. In spring,the terminal bud started expanding before the ‘whorl’buds which preceded the ‘interwhorl’ stem buds;completion of the flush of growth, determined by the end ofleaf expansion, occurred in the reverse order, ‘interwhorl’> ‘whorl’ > terminal. Similarly bud expansionstarted at the top of the crown and progressed downwards, andthe first shoots to complete their flush were at the bottomof the crown. Approximately 60% of the buds on each shoot beganexpanding in spring but only about half of these formed branches.Bud abscission began in May and by Sep. 45% of buds originallypresent had abscised. Most of-the buds that did not abscisewere the small buds at the base of the shoot that were not originallyassociated with a leaf. Approximately 42% of ‘whorl’buds and 28% of MnterwhorP stem buds formed branches. ‘Whorl’branches were approx. 60% longer that ‘interwhorl’stem branches; buds on the lower surface of the shoot producedlonger branches than those on the upper surface. The implicationsof the results for the development of crown form and selectionof superior oak are discussed. Quercus petraea, oak, buds, branches, crown form     

Sugar transport together with environmental conditions controls time lags between xylem and stem diameter changes

In the present study the seasonal patterns of time lags between diurnal xylem and whole stem diameter variations at the top and at the base of two Scots pine trees (Pinus sylvestris L.) were compared. The diameter variations were measured during the summers of 2001 and 2002. Time lags were determined using the cross‐correlation method. The lags were found to vary in time according to the different stages of growth. At the top the xylem lagged behind the whole stem between the beginning of stem growth and the end of shoot growth in both years. In 2001 the time lags at the base showed a similar behaviour during stem growth. That kind of seasonal pattern of the time lags would result from the changes in the sink strength due to changing growth rate at different parts of the tree and the differences in the annual rhythm of growth and water availability in the soil (based on precipitation measurements) between the years 2001 and 2002 were reflected in the patterns. The time lags of shrinking and swelling periods during high and low photosynthetic activity (measured using a shoot chamber) were also compared. It was found, for example, that in 2001 in the middle of the growing season at the top of the tree the whole stem lagged on average 15 min more behind the xylem on the days of high photosynthetic activity than on the days of low or moderate. These results show for the first time that the transportation of carbohydrates and variable sink activity could be detected during the growing season in field conditions using stem and xylem diameter variation measurements. Furthermore, these results provide evidence of the pressure gradient‐driven flow also in the phloem of gymnosperms.     

Rapid Oscillations in Plant Water Potential Measured with a Stem Psychrometer

Oscillations in water potential were measured for the firsttime in the plant stem. The phase of the oscillations in waterpotential (measured in Brussels sprout) lagged behind that oftranspiration by less than 5 min. The cycles ranged from 22to 48 min. The results support the view that oscillations inwater potential in the plant stem and interconnecting xylemtissues would synchronize oscillations in transpiration andstomatal opening throughout the plant. Brassica oleraceae L., Brussels sprout, oscillations, stem water potential, transpiration, stem psychrometer     

Stomatal oscillations in orange trees under natural climatic conditions

BACKGROUND AND AIMS: Stomatal oscillations have been reported in many plant species, but they are usually induced by sudden step changes in the environment when plants are grown under constant conditions. This study shows that in navel orange trees (Citrus sinensis) pronounced stomatal oscillations occur and persist under natural climatic conditions. METHODS: Oscillations in stomatal conductance were measured, and related to simultaneous measurements of leaf water potential, and flow rate of sap in the stems of young, potted plants. Cycling was also observed in soil-grown, mature orchard trees, as indicated by sap flow in stem and branches. KEY RESULTS: Oscillations in stomatal conductance were caused by the rapid propagation and synchronization of changes in xylem water potential throughout the tree, without rapid changes in atmospheric conditions. CONCLUSIONS: The results show marked stomatal oscillations persisting under natural climatic conditions and underscore the need to discover why this phenomenon is so pronounced in orange trees.     

Riverine networks constrain β‐diversity patterns among fish assemblages in a large Neotropical river

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Involvement of indole-3-acetic acid in the circadian growth of the first internode of Arabidopsis

The extension rate of the first inflorescence node of Arabidopsis was measured during light/dark or continuous light exposure and was found to exhibit oscillations which showed a circadian rhythmicity. Decapitation induced a strong inhibition of stem extension. Subsequent application of IAA restored growth and the associated extension–rate oscillations. In addition, IAA treatments, after decapitation, re-established the circadian rhythmicity visible in the intact plants during free run. This indicates that the upper zone of the inflorescence has a major influence on the extension rate of floral stems and implies a role for auxin. Application of N-(1-naphthyl)phthalamic acid, an IAA transport inhibitor, to an intact floral stem inhibited growth and the rhythmicity in the extension rate oscillations, indicating that IAA polar transport may play a role in the dynamics of stem elongation. Furthermore, IAA-aspartate application, after decapitation, did not restore growth and rhythmicity. Nevertheless, biochemical analysis of IAA and IAA-aspartate demonstrated circadian fluctuations of the endogenous levels of both compounds. These observations suggest that IAA metabolism is an essential factor in the regulation of the circadian growth rhythm of Arabidopsis floral stems. Received: 21 September 1998 / Accepted: 23 January 1999     

Multiple resonance damping or how do trees escape dangerously large oscillations?

To further understand the mechanics of trees under dynamic loads, we recorded damped oscillations of a Douglas fir (Pseudotsuga menziesii) tree and of its stem without branches. Eigenfrequencies of the branches were calculated and compared to the oscillation frequency of the intact tree. The term eigenfrequency is used here to characterize the calculated resonance frequency of a branch fixed at the proximal end to a solid support. All large branches had nearly the same frequency as the tree. This property is a prerequisite for the distribution of mechanical energy between stem and branches and leads to an enhanced efficiency of damping. We propose that trees constitute systems of coupled oscillators tuned to allow optimal energy dissipation.     

The Effects of Temperature and Other Variables on the Contribution of Lateral Branches to Yield in Phaseolus vulgaris L.

At low temperatures (15/15 °C day/night) in controlled environmentsthe growth of lateral branches at the cotyledonary node ofPhaseolusvulgarts L is suppressed The suppression can be overcome byraising the temperature of the buds by approximately 6 °Cwith small electric heaters In order to test the practical significanceof the induction of branching for yields of the commercial crop,seedlings were raised in contrasted regimes and then transplantedto the field The effects of pretreatment regime on final yieldwere small, changes in the yield from axillary branches tendedto be balanced by compensatory changes in the yield from themain stem In another field experiment, synthetic growth substanceswere applied in order to suppress or enhance branching Changesin the amount of yield carned on branches were again offsetby compensatory changes in the yield from the main stem Compensatoryeffects between branches and main stem were also found in avariety trial However, in an experiment on a single cultivarand various levels of N fertilizer, compensatory effects werenot found, here, branch and main stem yields were positively,rather than negatively, correlated These results are discussedin relation to the intrinsic factors that govern yield in Pvulgaris Phaseolus vulgaris L, dwarf bean, axillary branches, correlative inhibition, temperature, growth substances, plant density, yield     

Model simulations of continuous rotifer cultures

Derived from the Monod-model and regulating principles a regulation model of the rotifer development in chemostats was developed. The model was validated in continuous cultures of Brachionus angularis both in steady-states, when undisturbed, and in transient-states after perturbations by step changes of dilution rate or input substrate concentration. Simulations of the simple model monotonically approached steady-states, but cultures show overshoots and damped oscillations before reaching this state. After introducing time-lags into the model it depends on the size of the time lag if model rotifer densities reach stable steady-state values (at low time lags) or stable limit cycles with periodic oscillations (at high time lags). At even higher time lags chaotic conditions occur in the model with final extinction of the rotifers.     

Whole-tree level water balance and its implications on stomatal oscillations in orange trees [Citrus sinensis (L.) Osbeck] under natural climatic conditions

Sustained cyclic oscillations in stomatal conductance, leaf water potential, and sap flow were observed in young orange trees growing under natural conditions. The oscillations had an average period of approximately 70 min. Water uptake by the roots and loss by the leaves was characterized by large time lags which led to imbalances between water supply and demand in the leaves. The bulk of the lag in response between stomatal movements and the upstream water balance resided downstream of the branch, with branch level sap flow lagging behind the stomatal conductance by approximately 20 min while the stem sap flow had a much shorter time lag of only 5 min behind the branch sap flow. This imbalance between water uptake and loss caused transient changes in internal water deficits which were closely correlated to the dynamics of the leaf water potential. The hydraulic resistance of the whole tree fluctuated throughout the day, suggesting transient changes in the efficiency of water supply to the leaves. A simple whole-tree water balance model was applied to describe the dynamics of water transport in the young orange trees, and typical values of the hydraulic parameters of the transpiration stream were estimated. In addition to the hydro-passive stomatal movements, whole-tree water balance appears to be an important factor in the generation of stomatal oscillations.     

Bud and crown architecture of white spruce and black spruce

Needle primordia in buds and branch lengths were assessed in the crown of a plantation-grown white spruce tree. There was a gradation in needle primordia in buds in branches within the crown. The largest number of primordia was in the terminal bud of the leading main stem shoot, with the number in first-order whorl lateral shoot terminal buds decreasing from whorl 1 to whorl 4, below which buds contained a similar small number of primordia (about one-third as many as in the terminal shoot). Previous year's shoot elongation followed a similar pattern (i.e., elongation of whorl branches was greater closer to the top of the tree and elongation in the fourth through ninth whorls was about one-third that of the main stem leader). Higher order branches within whorls had within-branch gradation in shoot elongation and number of needle primordia, with older branches having as few as 16–30 primordia in buds and 3–4 cm elongation for high-order branches on older main stem whorls. There were strong correlations between the number of primordia in branch terminal buds and branch length/diameter and bud length/diameter/volume. In both black spruce and white spruce, there were strong correlations of number of needle primordia in main stem leader terminal buds with number of needle primordia in terminal buds of first and second whorl leaders.     

Wind-induced stresses in cherry trees: evidence against the hypothesis of constant stress levels

We calculated the wind-induced bending moments and stresses generated in the stems of five Prunus serotina conspecifics differing in height and canopy shape and size (based on detailed measurements of stem projected area and location with respect to ground level) to test the hypothesis that wind-loads generate uniform and constant stress levels along the lengths of tree twigs, branches, and trunks. These calculations were performed using five different wind speed profiles to evaluate the relative importance of the shape of wind speed profiles versus the ’geometry’ of tree shape on stem stress distributions and magnitudes. Additionally, we evaluated the effect of absolute tree size and stem taper on wind- induced stresses by scaling the size of smaller conspecifics to the absolute height of the largest of the five trees yet retaining the original stem proportions (i.e., diameter relative to stem length) for each plant. Finally, we also determined how the factor of safety for wind-loading (i.e., the quotient of stem yield stress and wind-load stress) changed as a function of tree size (and, presumably, age). Our results indicate that wind-load stress levels (1) vary along stem length even for the same wind speed profile and the same maximum wind speed; (2) would increase to dangerous levels with increasing tree height if it were not for ontogenetic changes in stem taper and canopy shape that reduce stress intensities to manageable levels; (3) tend to be more dependent on stem taper and canopy shape and size than on the shape of the wind speed profile; and (4) the factor of safety against wind-induced mechanical failure decreases as trees get larger, but varies along the length of large trees such that preferential stem failure is likely and functionally adaptive. We thus (1) reject the hypothesis of constant wind-induced stress levels; (2) support the view that size-dependent changes in stem taper are required to maintain wind-load mechanical reliability; and (3) suggest that certain portions of mature trees are ’designed’ to fail under high winds speeds, thereby reducing drag and the bending moments and stresses experienced by trunks. Received: 24 May 1999 / Accepted: 8 October 1999     

Temporal patterns of water flux in trees and lianas in a Panamanian moist forest

Using constant heat sap flow sensors, xylem water fluxes in ten tree species and two liana species were monitored for 5–10 days during the beginning of the wet season in May, 1993. For a subset of the trees, a branch was also monitored at the top of the crown for 5 days. Xylem flux (J _S) was related diurnally in all plants to vapor pressure deficit (D) measured within the upper-third of the canopy, and to incoming shortwave radiation R _S above the canopy. Cross-correlation analysis was used to estimate time lags between diurnal patterns of J _S and D or R _S, and between J _S in stems and branches. The maximum correlation coefficient from cross-correlation of J _S with R _S (range=0.57–0.92) was often higher than the maximum of J _S with D (range=0.43–0.89), indicating that diurnal J _S was more dependent on R _S than D. Time lags (lag corresponding to maximum correlation) of J _S at stem-base with D was shorter (0–45 min) than with radiation (5–115 min), highly variable within a species, and uncorrelated to the height or exposure of tree crowns or liana in the canopy. On a stand level, not accounting for the diel lag between stem sap flux and canopy flux resulted in errors in estimated canopy transpiration of up to 30%. Received: 19 October 1998 / Accepted: 8 June 1999     

High frequency spikes in long period blood cell oscillations

Several hematological diseases are characterised by oscillations of various blood cell populations. Two of these are a variant of chronic myelogenous leukemia (CML) and cyclical neutropenia (CN). These oscillations typically have long periods ranging from 20 to 60 days, despite the fact that the stem cell cycling time is thought to be of the order of 2–3 days. Clinical data from humans and laboratory data from the grey collie animal model of CN is suggestive of the idea that these long period oscillations may also contain higher frequency spiky oscillations. We show how such oscillations can be understood in the context of slow periodic stem cell oscillations, by analysing a two component differential-delay equation model of stem cell and neutrophil populations.For Karl Hadeler, on his 70th birthday, leader, teacher, colleague and friend.     

CRENATICAULIS,A NEW GENUS OF DEVONIAN PLANTS ALLIED TO ZOSTEROPHYLLUM,AND ITS BEARING ON THE CLASSIFICATION OF EARLY LAND PLANTS

The name Crenaticaulis verruculosus is proposed for slender, pseudomonopodially and dichoto-mously branching plants that bore opposite to subopposite sporangia along the stem and two rows of prominent, multicellular teeth on opposite sides of the terete axis. Epidermal cells were either narrow and elongate parallel to the stem or short with a papillate outer tangential wall. Axillary tubercles were present on one side of the stem near lateral branches. Some tubercles bore remnants of branches. Sporangia dehisced along their distal margins into two unequal halves. No spores were found. Occasional short lengths of stem were petrified by iron pyrite. Sections revealed a cortex consisting of four to six rows of thick-walled cells and a xylem strand. The strand was elliptical in transverse section and maturation was exarch. Tracheids were chiefly scalariform. It is suggested that the so-called axillary tubercles, known in several plants of Devonian age, were scars of rhizophores like those in the modern genus Selaginella. The plant is referred to the subdivision Zosterophyllophytina.     

Existing branches correlatively inhibit further branching in Trifolium repens: possible mechanisms

In Trifolium repens removal of any number of existing branches distal to a nodal root stimulates development of axillary buds further along the stem such that the complement of branches distal to a nodal root remains constant. This study aimed to assess possible mechanisms by which existing branches correlatively inhibit the outgrowth of axillary buds distal to them. Treatments were applied to basal branches to evaluate the roles of three postulated inhibitory mechanisms: (I) the transport of a phloem-mobile inhibitory feedback signal from branches into the main stem; (II) the polar flow of auxin from branches into the main stem acting to limit further branch development; or (III) the basal branches functioning as sinks for a net root-derived stimulatory signal (NRS). Results showed that transport of auxin, or of a non-auxin phloem-mobile signal, from basal branches did not influence regulation of correlative inhibition and were consistent with the possibility that the intra-plant distribution of NRS could be involved in the correlative inhibition of distal buds by basal branches. This study supports existing evidence that regulation of branching in T. repens is dominated by a root-derived stimulatory signal, initially distributed via the xylem, the characterization of which will progress the generic understanding of branching regulation.