Chloroplast Distribution in the Stems of 23 Eucalypt Species
Abstract
:1. Introduction
- CAM plants (stem succulents, often with cladodes).
- Stem photosynthesis—green stems with epidermis (no or delayed periderm development) and a high stomatal density [9], with some species having a well-developed palisade layer (e.g., [10]). They are mainly herbaceous plants but not exclusively and include desert shrubs [10] and early successional legumes (references in [11]).
- Corticular or bark photosynthesis in woody plants after periderm development. Ávila et al. [9] indicate that this type of structure should be termed ‘cortical photosynthesis’. In stems of trees and shrubs, the zone between the periderm and the secondary phloem is often referred to as ‘cortex’. However, the cortex can be defined as a primary ground tissue between the epidermis and vascular bundles in a stem [12]. Thus, ‘secondary cortex’ [13] and ‘pseudo-cortex’ [14] have also been used for this zone. Rosell [15,16] considers that these cells are initially derived from the shoot apical meristem but then divide radially, thus are primary in origin (not derived from the vascular or cork cambia). Anatomical aspects of corticular photosynthesis have been more commonly investigated in small diameter/young stems (Table 1). Chloroplasts have been recorded in current year shoots to trunks 60 cm in diameter. In the few studies where both small and large diameters were examined, chloroplasts were found at greater depths in the smaller diameters (e.g., [17]). This may be related to light being conducted in the axial system of vessels, tracheids, and fibres (e.g., [18]). As could be expected, the density of chloroplasts decreases as phellem thickness increases (e.g., [3]).
- Wood or woody tissue photosynthesis usually occurs in the xylem ray parenchyma cells and can also extend into the pith in small diameter stems.
- i)
- given the findings of the studies in Table 1, chloroplasts would be found in all the small diameter branches as well as large diameter branches and trunks of smooth barked stems,
- ii)
- given the wide taxonomic diversity of species examined, different chloroplast arrangements would be recorded,
- iii)
- given that light appears to be conducted more effectively in small diameter stems, chloroplasts would be found at greater depth in smaller diameter stems than larger diameter ones,
- iv)
- the youngest stems (no periderm) would have stomata but they would be at a low density compared to adjacent leaves, and
- v)
- given that the species examined had thin phellem (provides little protection), the bark inside this layer would be optimised for defence and mechanical support rather than photosynthesis.
2. Results
2.1. External Morphology
2.2. Small Diameter Stems Without Periderm
2.3. Large Diameter Stems
3. Discussion
3.1. General Distribution of Chloroplasts
3.2. Differences in Chloroplast Distribution Between Species
3.3. Chloroplast Distribution in Small vs Large Diameter Stems
3.4. Stem and Leaf Stomatal Densities
3.5. Bark Functional Trade-Offs
3.6. Phellem Thickness and Chloroplast Development
3.7. Conclusions
4. Materials and Methods
4.1. Materials
4.2. Microscopy
4.3. Light Transmission
4.4. Maximum Chloroplast Depth
4.5. Stomatal Density in Stems and Leaves
Author Contributions
Funding
Acknowledgments
Conflicts of Interest
References
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Author | Date | Species | Materials Examined |
---|---|---|---|
Scott [38] | 1907 | 30 woody species | Young shoots and inflorescence axis |
Sokolov [33] | 1953 | 100 angiosperm spp., 16 gymnosperm spp. | |
Pearson & Lawrence [39] | 1958 | Populus tremuloides | 19 yr old, 16 cm diameter at 2 m above ground level |
Gómez-Vazquez & Engleman [40] | 1984 | Bursera longipes, B. copallifera | Trees 8 and 5 m high, respectively (no diameter given) |
Kauppi [41] | 1991 | Betula pendula, B. pubescens | 0, 1, 3, 4, 5, 10 and 20 yr old branches |
van Cleve et al. [42] | 1993 | Populus × canadensis | 2–3 yr old twigs, 1–2 cm diameter |
Pilarski [43] | 1999 | Syringa vulgaris | Current year, 1 and 3 yr old stems |
Pilarski & Tokarz [17] | 2006 | Fagus sylvatica | 2 and 6 yr old stems and trunk (3.4, 6.9 and 600 mm diameter, respectively) |
Dima et al. [34] | 2006 | 20 species | 2 yr old twigs |
Berveiller et al. [44] | 2007 | 9 tree species | Current year stems, 2-5 mm diameter |
Filippou et al. [5] | 2007 | Olea europaea | 1–30 yr old stems |
Rentzou & Psaras [6] | 2008 | 3 Mediterranean species | 1–2 yr old and 2–5 yr old twigs |
Kotina et al. [45] | 2012 | 14 species Apiaceae | Branch tips to thicker stems with mature bark |
Schmitz et al. [46] | 2012 | 13 Australian mangrove species | Mainly 6 mm diameter (range 2.8–14 mm diameter) |
Cocoletzi et al. [47] | 2013 | Myriocarpa longipes, Urera glabriuscula | 10 yr old trees |
Kocurek et al. [48] | 2015 | Clusia multiflora, C. rosea | 2.0–2.5 cm diameter at 1 m above ground level |
Wittmann & Pfanz [49] | 2016 | 5 woody species | Current year shoots, 3–5 mm diameter |
Kotina et al. [14] | 2017 | Adansonia digitata | Young twigs to stems 12–17 cm diameter |
Blagitz et al. [50] | 2019 | Monquinastrum polymorphumZanithoxylum rhoifolium | Large diameter stems |
Species | Leaf (Adaxial) | Leaf (Abaxial) | Stem |
---|---|---|---|
Corymbia citriodora | 267 (± 28) | 320 (±39) | 2.3 (± 1.2) |
Corymbia ficifolia | 0 (± 0) | 209 (±39) | 1.6 (± 0.8) |
Eucalyptus cladocalyx | 0 (± 0) | 279 (±27) | 1.9 (± 0.8) |
Eucalyptus melliodora | 184 (± 31) | 194 (±60) | 4.1 (± 2.1) |
Eucalyptus sideroxylon | 134 (± 17) | 169 (±22) | 1.3 (± 1.1) |
Eucalyptus torquata | 105 (± 24) | 124 (±26) | 2.2 (± 1.6) |
Genus/Species | State/Territory | Subgenus-Section | Bark Type | Stem Diameter | Phellem Thickness (μm) | Chlorenchyma Bright Band (μm) | Chlorenchyma Maximum Depth (μm) |
---|---|---|---|---|---|---|---|
Angophora | |||||||
A. hispida (Sm.) Blaxell | NSW | Rough to small branches, fibrous | S (2 mm) S (6–8 mm) M (13–15 mm) | - 30–70 30–60 | 100–200 150–200 200 | 600–1000 800–1000 1700–1800 | |
Corymbia | |||||||
C. citriodora (Hook.) K.D. Hill & L.A.S. Johnson | Qld | Blakella | Smooth throughout—gum | S (1.5–3 mm) S/M (5–11 mm) L (10–40 cm) | - 50–70 20–90 | 150–200 100–150 100–330 | 1000–1400 500–1500 700–1300 |
C. maculata (Hook.) K.D. Hill & L.A.S. Johnson | NSW, Qld, Vic | Blakella | Smooth throughout | S (3–6 mm) M (11–14 mm) L (30–50 cm) | 0–60 30–40 30–50 | 150–200 120–150 120–200 | 1000 1400–1500 900–1800 |
C. ficifolia (F. Muell.) K.D. Hill & L.A.S. Johnson | WA | Corymbia | Rough to small branches, fibrous | S (3–6 mm) M (14–15 mm) M (26 mm) | - 50–100 50–80 | 220–400 250–300 300 | 1200–2000 1800–2100 1800–2100 |
C. eximia (Schauer) K.D. Hill & L.A.S. Johnson | NSW | Blakella | Rough to small branches; tessellated, flaky | S (2–7 mm) M (10–30 mm) L (7–10 cm) | - 50–100 100 | 200–400 100–400 100–120 | 1100–2000 1000–3000 1100–1400 |
Eucalyptus | |||||||
E. erythrocorys F. Muell. | WA | Eudesmia | Smooth; can have rough bark on lower trunk | L (4–5 cm) L (15–20 cm) | 50–80 60–100 | 80–100 100 | 2300–3000 2500–2800 |
E. macrorhyncha F. Muell. ex Benth. | NSW, Vic, SA | Eucalyptus | Rough to small branches (stringybark) | S (2 mm) M (8–18 mm) | - 20–100 | 120–150 120–220 | 900–1000 1000–3000 |
E. rossii R.T. Baker & H. G. Sm. | NSW | Eucalyptus | Smooth (scribbly gum) | S (2 mm) M (11–20 mm) L (10–20 cm) | - 30–80 30–100 | 120–150 200–400 150–280 | 900–1000 1500–2000 1000–1300 |
E. leucoxylon F. Muell. | SA, Vic | Symphyomyrtus—Adnataria | Smooth throughout, sometimes with rough box-type at base | S (1.5–4.5 mm) S (7 mm) L (17 cm) | - 20 80–90 | 170–200 150 150–300 | 800–2000 1300 700–1100 |
E. melliodora A. Cunn. ex Schauer | NSW, Vic, Qld | Symphyomyrtus—Adnataria | Rough box-type at base, smooth higher | S (2.5–3 mm) S (6–8 mm) L (50–80 cm) | - 30–50 50–100 | 150–200 150–200 100–150 | 1100–1500 700–1400 700–800 |
E. sideroxylon A. Cunn. ex Woolls | NSW, Vic, Qld | Symphyomyrtus—Adnataria | Rough throughout (ironbark) | S (3–8 mm) M (10–12 mm) M (23–26 mm) | - 30–70 40–60 | 200 150–250 100–200 | 1000–1900 1800–2200 800–2300 |
E. albens Benth. | NSW, Vic, Qld, SA | Symphyomyrtus—Adnataria | Rough on trunk and large branches (box-type) | S (2–4 mm) M (12–28 mm) | - 50–80 | 250–400 150–350 | 700–1900 900–1500 |
E. caesia Benth. | WA | Symphyomyrtus—Bisectae | Minnirichi on trunk and branches | *1 | |||
E. macrocarpa Hook. | WA | Symphyomyrtus—Bisectae | Smooth | S/M (3–15 mm) L (8–10 cm) | - 40–50 | 200–500 100–200 | 2000–3200 800–1100 |
E. kruseana F. Muell. | WA | Symphyomyrtus—Glandulosae | Rough but thin on trunk base, smooth above | S (3–8 mm) M (15–40 mm) | - 100 | 200–300 50–100 | 1000–2600 500–1500 |
E. torquata Luehm. | WA | Symphyomyrtus—Dumaria | Rough, hard, shortly fibrous to almost tessellated | S (3–8 mm) M (15–30 mm) | - 100–300 | 250–400 100–300 | 1000–3500 500–1400 |
E. blakelyi Maiden | NSW, Vic, Qld | Symphyomyrtus—Exsertaria | Smooth throughout (gum) | S (3–7 mm) M (10–40 mm) L (20–37 cm) | - 20–60 30–100 | 200–300 100–300 100–200 | 1200–1600 1500–2000 300–700 |
E. camaldulensis Dehnh. | Most states | Symphyomyrtus—Exsertaria | smooth | S (3–6 mm) M (8–25 mm) L (6–80 cm) | - 30–50 30–70 | 150–200 100–200 100–200 | 900–1800 500–800 500–800 |
E. globulus Labill. | Tas, Vic | Symphyomyrtus—Maidenaria | Smooth apart from the base with persistent slabs (gum) | S (3–8 mm) M (18–20 mm) L (50–60 cm) | - 30–40 30–80 | 200–300 150–200 150–300 | 1300–2800 1000–1800 800–1200 |
E. cinerea F. Muell. ex Benth. | NSW | Symphyomyrtus—Maidenaria | Thick, furrowed, fibrous to small branches | S (2–6 mm) M (10–20 mm) | - 10–80 | 150–200 100–250 | 900–2700 1500–3500 |
E. nicholii Maiden & Blakely | NSW | Symphyomyrtus—Maidenaria | Thick, fibrous to small branches (peppermint) | S (3–4 mm) M (8–18 mm) | - 20–50 | 150–200 200–350 | 1300–1900 1700–2300 |
E. scoparia Maiden | Qld | Symphyomyrtus—Maidenaria | Smooth—gum | S (3–8 mm) L (4–26 cm) | - 30–50 | 150–300 150–400 | 1400–2800 900–1300 |
E. cladocalyx F. Muell. | SA | Symphyomyrtus—Sejunctae | Smooth—gum | S (2–8 mm) L (24–60 cm) | 0–60 40–70 | 180–300 80–150 | 800–3900 600–800 |
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Burrows, G.E.; Connor, C. Chloroplast Distribution in the Stems of 23 Eucalypt Species. Plants 2020, 9, 1814. https://0-doi-org.brum.beds.ac.uk/10.3390/plants9121814
Burrows GE, Connor C. Chloroplast Distribution in the Stems of 23 Eucalypt Species. Plants. 2020; 9(12):1814. https://0-doi-org.brum.beds.ac.uk/10.3390/plants9121814
Chicago/Turabian StyleBurrows, Geoffrey E., and Celia Connor. 2020. "Chloroplast Distribution in the Stems of 23 Eucalypt Species" Plants 9, no. 12: 1814. https://0-doi-org.brum.beds.ac.uk/10.3390/plants9121814