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Protea Night-Life: Ecological and Commercial Significance for Soil Hydraulics 1

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Protea Night-Life: Ecological and Commercial Significance for Soil Hydraulics 1 Protea night-life: Ecological and commercial significance for soil hydraulics 1. Heidi-J Hawkins A,C, Hans Hettasch B and Michael D Cramer A ADepartment of Botany, University of Cape Town, Private Bag X1, Rondebosch, 7701, South Africa; BArnelia Farms, P.O. Box 192, Hopefield, 7355, South Africa CCorresponding author Tel: +27 21 650 2442 Fax: +27 21 650 4041 Email: [email protected] 1 Submitted Feb 2009 in detailed format to Functional Plant Biology as ‘Hydraulic redistribution by Protea ‘Sylvia’ (Proteaceae) facilitates soil water recharge and water acquisition by an understorey grass and shrub’. Please use the latter article to cite information . 1 Introduction During the day , if plant stomata are open, water moves along the water potential (ψw) gradient from soil to leaf to atmosphere via the xylem , i.e . transpiration occurs (Fig. 1a) . At night , or whenever stomata close , that gradient is removed ... However,However,However, asas longlong asas plantplant rootsroots spanspan wetwet (usually(usually deeper)deeper) and drier (usually shallow) soil areas, another water gradient exists , i.e. between wet ter and drier soil layers. In this case, w ater moves along the ψw gradient from wet soil toto thethe root,root, along the root xylem andand outout intointo drier soil , resulting in soil wetting (Fig. 1b). This passive process is called hydraulic redistribution and has been documented for more than 50 taxa includingincluding trees, shrubs and grasses , particularly in summer -drought Mediterranean areas, but also in deserts , tropical forests (Jackson et al. 2000) and crop plants ( Sekiya and Yano 2004 ). Hydraulic redistribution is thought to be important forfor soilsoil waterwater recharge and facilitating nutrientnutrient uptake in water -limite d and summer -drought ecosystems, e.g. Great Basin of the western United States ; Mediterranean -type ecosystems such as the Australian kwongan, the Californian chaparral and the Cape Florist icic RegionRegion (CFR) of South Africa . However,However, the ecological and commercial importance of hydraulic redistribution has not been well -demonstrate d in the field. South African Proteaceae are often the largest shrubs in the CFR (2 up to 10 m, Rebelo 2001) and havehave been shown to continue to transpire throughout thethe summersummer droughtdrought in the west ofof thethe CFRCFR whilewhile other shallower -rooted plants cease to tr anspire (van der Heyden and Lewis, 1988). Not only do ProteaceaeProteaceae continue to transpire throughout summer,summer, but Proteaceae grow predominantly duringduring thethe hot,hot, dry summer months (Rebelo 2001).2001). Roots are dimorphic with deep ‘sinker ’ roots and shallow lateral ro ots (Mortimer et al. 2003) si milar to Australian Proteaceae (Pate et al. 1995). Rooting depths for shrub -forms of SA Proteaceae are known to reach at least 3.6 m (Higgins et al. 1987) but may extend deeper depending on the water table . This implies that the deeper root ed Proteaceae have access to deep water reserves which sustain photosynthesis, transpiration and growth through the summer. Fig. 1. Soil water moves along water potential OurOur preliminary data using soil moisture probes gradients. In the day the gradient is usually from soil showed that soil moisture in summer near lateral toto plantplant toto atmosphereatmosphere (a), during thethe nightnight oror whewhenn roots (0.2 -0.4 m d epth) of Protea ‘Sylvia’ was stomatastomata areare closed and the gradient to atmosphere iiss indeed higher than slightly deep er in the soil profile removed, wat er can move from wet to drier soil in a (e.g. 0.5 m, Fig. 2 ) while being similar to deeper process called hydraulic redistribution (b). Pictures soil layers ( 1.0 m, Fig. 2) suggesting that hydraulic modified with permission courtesy of FC Meinzer, redistributionredistribution toto dry upper soil layers was occurrioccurring.ng. USDA Forest Servic e, Forest Science Lab, Corvalis, Therefore, we hypothesize d that Proteaceae are able OR, 2003. to hydraulic ally redistribute water from deeper 2 moist soil layers to upper drier soil layers where nutrients occur. Redistributed water may aid in improving the water and nutrient status of the proteas, but it may also positively influence surrounding plants (Dawson 1993) and thus may be an important component maintaining diversity in these systems. In addition, if commercially grown Proteaceae are encouraged to have deep root systems via ‘deep’ irrigation versus frequent shallow irrigation events, these crops may be made more tolerant to drought stress. Fig. 2 Soil moisture profiles adjacent to Protea ‘Sylvia’ plants grown on aeolian sands at Arnelia Farms, Hopefield, South Africa, before supply of heavy water label. Values are means ± standard error (n=4). Brief materials and methods Our field experiment was accomplished by sinking PVC plumbing pipes 1.2 m into the soil profile next to farmed five-year old Protea ‘Sylvia’ plants. Plants were irrigated via these pipes to simulate deep water supply and to encourage root growth near the pipe. After ca. 2 months, plants were either supplied with 25 L labelled water overnight or not. Labelled water, or so-called heavy water was characterized by the ratio of heavy 3 hydrogens (2H) to light hydrogens (1H) as δ2H (measured in ‰). Any increase in δ2H of the upper soil overnight was interpreted as evidence of hydraulic redistribution or hydraulic lift. Likewise, an increase in δ2H of shallow-rooted plants ( Leysera gnapholodes , Asteraceae) under the protea was interpreted as evidence of water parasitism by the shrub, since the shrub had no access to water at depth. Lithium, an analogue of potassium that is not metabolised by plants and thus may be traced, was placed around plants receiving labelled water to determine whether a short-term supply of water at depth, and subsequent hydraulic redistribution, could contribute to nutrient uptake. A pot experiment with Sylvia and the shallow-rooted Fynbos grass, Cyanodon dactylon (Poaceae), was used to support field findings. In addition, an irrigation experiment tested the practical implications of hydraulic redistribution on sandy soils. In this experiment, Sylvia received 90 L water week -1, supplied via surface drippers (shallow irrigation), half surface drippers and half via a 1.2 m deep pipe (half shallow/half deep irrigation) or via the 1.2 m pipe only (deep irrigation). Deep irrigation was meant to simulate less frequent, high volume irrigation. Plants receiving deep irrigation were acclimatized to this irrigation over a few months to allow for root growth near the pipe. Results and Discussion It was found that the large Sylvia plants growing in aeolian sands in the so-called Strandveld Fynbos (Arnelia Farms, Hopefield, on the west coast of the CFR, 33 °2’49’’S, 18 °20’30’’E) took up the labelled water at a depth of 1.2 m as was evident from the label found in Sylvia stems (Fig. 3a). This confirmed that Sylvias had root access to the water supplied at depth, as was hoped. More importantly, Sylvia redistributed labelled water from 1.2 m depth to the 0.2 to 0.4 m soil layer, as indicated by increased δ2H of soil water from -24.5 ± 0.7‰ to - 8.0 ± 3.0‰ (Fig. 3b) and a 52% increase in soil water from 0.48% to 0.89% moisture content. Linear mixing models attributed 0.68 ± 0.05% of the water in the upper soil layer to the label, or 8.5 ± 0.9% when using Sylvia plants as the label source. This is a rather small redistribution of water and may be linked to the short-term nature (overnight) of the experiment. The small amount of hydraulic redistribution was apparently insufficient to result in there being any difference in Li uptake between plants receiving 25 L heavy water label at depth and those receiving no additional water overnight. This suggests that hydraulic redistribution does not make a measurable contribution to ion uptake in the short-term. However, both treatments accumulated considerable amounts of Li over the three week period subsequent to watering, meaning that both treatments had access to sufficient soil moisture to allow the Li to diffuse to the roots. It is possible that plants in both treatments made use of hydraulically-lifted water from >2m depth (from trenches we know that Sylvia roots extend at least 2 m). Therefore, the amounts of additional water supplied at 1.2 m depth may have made no difference to the amount of Li in soil solution or to that taken up by the plant in this sandy soil. Snyder et al. (2008) found that hydraulically redistributed water did not increase the amount of 15 N acquired by the desert shrub Sarcobatus vermiculatus in dune sand. Both the present and the latter study were performed on sandy soils where hydraulic redistribution may be less important due to relatively less root-soil contact compared to clayey soils. Also matric forces (capillary and electrostatic) between water and sands are small due to the relatively large pore sizes of the sand. Relatively larger gravimetric forces may mean that hydraulically-lifted water quickly drains out of the upper sand layers. Water in stems of the shallow-rooted understorey shrub, L. gnaphalodes had similar δ2H values (338 ± 157‰) to stems of Sylvia, indicating water parasitism by L. gnaphalodes . Supporting this field data was pot 3 experiments where Sylvias redistributed ca . 17% of another type of water label (radioactive H2O) supplied, equating to 34 ± 1.2 mL plant -1. Also, the shallow-rooted local grass, Cyanodon dactylon (Poaceae), growing in the same pots as Sylvia were labelled, again indicting water ‘parasitism’. In the field, the two understory plants are likely dependent on periodic rainfall events during summer and/or reliant on water redistributed by large shrubs. Whatever the competitive interactions between protea and understorey plants during recruitment, once established, the persistence of grass and small shrubs such as the L.
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